强光下硫化氢通过促进光系统Ⅱ的活性来缓解水稻的光抑制

以水稻品种‘II优084’为材料,测定了强光胁迫下,水稻光合速率、叶绿素荧光快速诱导曲线(OJIP)以及O2ˉ·和H2O2在水稻叶片中积累的影响。结果表明强光胁迫下,水稻的净光合速率及气孔导度下降;光系统II(PSII)反应中心关闭的比例以及电子传递链中光系统II受体侧原初醌受体(QA)的还原程度增加;PSII反应中心电子传递的量子产额、能量以及传递到下游电子链的比率下降;光抑制下PSII的过剩能量向PSI的状态装换减少;自由基的产生增加。而施加作为硫化氢(H2S)供体的外源硫氢化钠(NaHS)后,上述影响PSII活性的指标的负变化被缓解,捕光天线复合体LHC通过在两个光系统之间的移动,来调节两个光系统的能量分配。强光下H2S处理能促进LHC离开PSII,与PSI结合,从而减少PSII分配的激发能,增加PSI分配的激发能,缓解了PSII的过度还原。以上结果表明外源H2S通过促进PSII的光合活性来缓解水稻光抑制伤害。     

NaCl胁迫加重强光胁迫下超大甜椒叶片的光系统II和光系统I的光抑制

快速叶绿素荧光动力学可以在无损情况下探知叶片光合机构的损伤程度, 快速叶绿素荧光测定和分析技术(JIP-test)将测量值转化为多种具有生物学意义的参数, 因而被广泛应用于植物光合机构对环境的响应机制研究。该文研究了超大甜椒(Capsicum annuum)幼苗在强光及不同NaCl浓度胁迫下的荧光响应情况。与单纯强光胁迫相比, NaCl胁迫引起了叶绿素荧光诱导曲线的明显改变, 光系统II (PSII)光抑制加重, 同时PSII反应中心和受体侧受到明显影响, 而且高NaCl浓度胁迫下PSII供体侧受伤害明显, 同时PSI反应中心活性(P700⁺)在盐胁迫下明显降低。这些结果表明, NaCl胁迫会增强强光对超大甜椒光系统的光抑制, 并且浓度越高抑制越明显, 但对PSI的抑制作用低于PSII。高NaCl浓度胁迫易对PSII供体侧造成破坏, 且PSI光抑制严重。     

2，3－环氧丙酸钾对水稻光合功能的改善

光呼吸抑制剂2,3－环氧丙酸钾对水稻光合功能具有明显的改善作用,不论在整体叶片水平还是在离体叶绿体条件下,均能提高光系统II（PSII）的活性和原初光能转化效率;提高叶绿体的PSII和光系统I（PSI）及全链（PSII＋PSI）的电子传递速率;同时它还可以提高镁离子（Mg^2+)和叶绿体膜蛋白磷酸化对两个光系统之间光能分配的调节能力,结果表明了2,3－环氧丙酸钾在水稻生产上有一定应用前景。     

2，3－环氧丙酸钾对水稻光合功能的改善

光呼吸抑制剂2,3－环氧丙酸钾对水稻光合功能具有明显的改善作用,不论在整体叶片水平还是在离体叶绿体条件下,均能提高光系统II（PSII）的活性和原初光能转化效率;提高叶绿体的PSII和光系统I（PSI）及全链（PSII＋PSI）的电子传递速率;同时它还可以提高镁离子（Mg^2 )和叶绿体膜蛋白磷酸化对两个光系统之间光能分配的调节能力,结果表明了2,3－环氧丙酸钾在水稻生产上有一定应用前景。     

外源葡萄糖对盐胁迫下山楂叶中光系统Ⅱ光化学活性的影响

研究盐胁迫下外源葡萄糖对山楂叶中光系统II（PSII）光化学活性影响的结果表明,盐胁迫下,浇灌外源葡萄糖可增加山楂幼苗叶中PSII最大光化学效率（Fv/Fm）、暗适应后PSII最大光化学效率（ΦPo）、捕获的激子将电子传递到电子传递链中QA-下游的其它电子受体的概率（Ψo）及反应中心吸收的光能用于电子传递的量子产额（ΦEo）,降低照光2ms时反应中心的关闭程度（Vj）和单位反应中心吸收的能量（ABS/RC）,提高电子转运效率（ETo/RC）,降低放氧复合体（OEC）受伤害的程度。     

短期生长环境光强骤增导致典型阴生植物三七光系统受损的机制

阴生植物突然暴露在强光下造成光损伤的情况时有发生, 但其对高光敏感的潜在机制尚不十分清楚。为阐明阴生植物无法在自然全光照环境下生存的相关机制, 该研究以典型阴生植物三七(Panax notoginseng)为材料, 将遮阴环境下(10%透光率)生长的植株转移到全日光环境下3天, 研究其相对叶绿素含量(SPAD值)、光合参数以及叶绿素荧光参数的变化。结果表明, 全光环境下三七光合日变化呈现“双峰”曲线特征, 且净光合速率在处理期间逐日降低。全日光下三七叶片SPAD值、水分利用率和光能利用率显著降低; 叶片光系统I (PSI)反应中心P700最大荧光信号、光系统II (PSII)电子传递速率、暗适应下PSII最大量子效率和光下PSII最大量子效率显著低于遮阴环境下的植株, 且至傍晚不能完全恢复。而参与调节性能量耗散的量子产量、PSI受体侧限制引起的非光化学量子产量、环式电子流则显著高于遮阴环境下的三七。此外, 生长环境光照强度骤增导致荧光诱导动力学曲线发生明显变化, 并显著升高了PSII供体侧和受体侧的荧光产量。当阴生植物三七突然暴露于全光环境下时, 强烈的光照会导致PSII供体侧的放氧复合体活性受损, 抑制受体侧的电子传递, 过度还原PSI的受体侧进而引发PSI光抑制。该研究结果揭示, 全日光导致的PSII不可逆损伤和PSI光抑制可能是典型阴生植物三七为什么不能在全日照光环境下存活的重要原因。     

外源Ca2＋对高温强光胁迫下灌浆期小麦叶片光合机构运转的影响

灌浆期叶面喷施10mmol·L-1 CaCl2对高温强光胁迫下小麦叶片光合电子传递、放氧速率、叶绿素荧光参数和D1蛋白的影响结果表明,Ca2＋预处理可保护D1蛋白,削弱其降解,提高光系统I（PSI）和光系统Ⅱ（PSⅡ）子传递速率、全链电子传递速率、净光合速率（Pn）、PSII最大光化学效率（Fv/Fm）、PSII实际光化学效率（ΦPSⅡ）和光化学猝灭（qp）,维持较低的Fo,最终导致小麦适应高温强光的能力提高。     

生长温度对烟草叶片环式电子传递活性的影响（英文）

高等植物的光合机构在环境胁迫条件下非常容易产生光抑制,环式电子传递在光合机构的光保护中发挥着重要的作用。但是,生长温度对环式电子传递的影响并不清楚。本研究测定了在24/18℃和32/26℃条件下生长40天的烟草(K326)叶片的气体交换、叶绿素荧光和P700氧化还原态的光响应曲线。结果表明,烟草叶片在两种生长温度下的的光合能力、光化学淬灭、非光化学淬灭和通过光系统II的电子传递速率(ETR II)均没有差异。但是,在强光条件下,生长在24/18℃的叶片比生长在32/26℃的具有更高的通过光系统I的电子传递速率(ETR I)和ETR I/ETR II比值。短时间的强光处理后,生长在24/18℃的叶片具有较高的光系统II最大量子产额(Fv/Fm),表明环式电子传递活性的上调有助于缓解生长在24/18℃的叶片光系统II受到的光损伤。综上所述,环式电子传递活性的增强是植物适应较低生长温度的重要策略。     

高温强光对温州蜜柑叶绿素荧光、D1蛋白和Deg1蛋白酶的影响及SA效应

以3年生温州蜜柑（Citrus unishiu Marc.）植株为试材,用叶绿素荧光分析、Western-blotting蛋白质印记技术及DAB（3,3'-二氨基联苯胺）显色法,研究了高温强光（38℃和1600 μmol?m-2?s-1）对叶片叶绿素荧光参数、PS（光系统）II反应中心D1蛋白和Deg1蛋白酶的影响和SA（水杨酸）的效应。结果表明,高温强光交互作用4 h后,叶片的初始荧光Fo升高,最大光能转化效率Fv/Fm、表观光合电子传递速率ETR及PSII的量子产额ΦPSII显著降低,在D1蛋白降解的同时,Deg1蛋白酶含量也下降,并伴有H2O2的积累。在高温强光下,外源的H2O2使叶绿素荧光动力学快相参数(Fi-Fo)/(Fp-Fo)值（反映PSII中QB非还原中心的数量）升高和I-P的斜率（反映PSII 活化中心还原态QA积累的值）下降,Fv/Fm、ETR、ΦPSII及D1蛋白和Deg1蛋白酶下降幅度增大;而外源的SA使这些参数下降幅度减小。这些结果说明,高温强光诱导H2O2的积累造成Deg1蛋白酶和光系统反应中心D1蛋白的降解,Deg1蛋白酶的减少也进一步限制了D1蛋白的周转,进而使温州蜜柑PSII反应中心遭到破坏,SA对光合机构光破坏有保护作用。     

外源6-BA对低温胁迫下茄子幼苗光合作用、叶绿素荧光参数及光能分配的影响

本文研究了外源6-BA对低温胁迫下茄子幼苗光合作用、叶绿素荧光参数和能量分配的影响。结果表明,外源6．BA显著增加了低温胁迫下茄子叶绿素含量、净光合速率（Pn）、蒸腾速率（t）、气孔导度（Gs）和胞间CO2浓度（c1）;同时外源6-BA明显提高了低温胁迫下茄子幼苗叶片的PSⅡ最大光化学效率（Fv/Fm）、PSⅡ潜在活性（R／Fo）、PSII天线转化效率（FvFm）、实际光化学效率（φpsⅡ）、光化学猝灭系数（g,）和光化学反应能量（P）,降低了非光化学猝灭系数（NPQ）、天线热耗散能量（D）,对非光化学反应耗散能量（E）无明显影响。表明外源6-BA处理通过促进低温胁迫下茄子幼苗光合作用,提高光合电子传递效率,从而保护光合系统,降低低温胁迫对植物的损伤。     

Enhanced function of non-photoinhibited photosystem II complexes upon PSII photoinhibition

Light induced photosystem (PS)II photoinhibition inactivates and irreversibly damages the reaction center protein(s) but the light harvesting complexes continue the collection of light energy. Here we addressed the consequences of such a situation on thylakoid light harvesting and electron transfer reactions. For this purpose, Arabidopsis thaliana leaves were subjected to investigation of the function and regulation of the photosynthetic machinery after a distinct portion of PSII centers had experienced photoinhibition in the presence and absence of Lincomycin (Lin), a commonly used agent to block the repair of damaged PSII centers. In the absence of Lin, photoinhibition increased the relative excitation of PSII and decreased NPQ, together enhancing the electron transfer from still functional PSII centers to PSI. In contrast, in the presence of Lin, PSII photoinhibition increased the relative excitation of PSI and led to strong oxidation of the electron transfer chain. We hypothesize that plants are able to minimize the detrimental effects of high-light illumination on PSII by modulating the energy and electron transfer, but lose such a capability if the repair cycle is arrested. It is further hypothesized that dynamic regulation of the LHCII system has a pivotal role in the control of excitation energy transfer upon PSII damage and repair cycle to maintain the photosynthesis safe and efficient.     

Mitochondrial alternative oxidase pathway protects plants against photoinhibition by alleviating inhibition of the repair of photodamaged PSII through preventing formation of reactive oxygen species in Rumex K-1 leaves

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.     

Light quality during growth of Tradescantia albiflora regulates photosystem stoichiometry, photosynthetic function and susceptibility to photoinhibition

Tradescantia albiflora (Kunth) was grown under two different light quality regimes of comparable light quantity: in red + far-red light absorbed mainly by photosystem I (PSI light) and yellow light absorbed mainly by photosystem II (PSII light). The composition, function and ultrastructure of chloroplasts, and photoinhibition of photosynthesis in the two types of leaves were compared. In contrast to regulation by light quantity (Chow et al. 1991. Physiol. Plant. 81: 175–182), light quality exerted an effect on the composition of pigment complexes, function and structure of chloroplasts in Tradescantia: PSII light-grown leaves had higher Chl a/b ratios, higher PSI concentrations, lower PSII/PSI reaction centre ratios and less extensive thylakoid stacking than PSI light-grown leaves. Light quality triggered modulations of chloroplast components, leading to a variation of photosynthetic characteristics. A larger proportion of primary quinone acceptor (Q_A) in PSI light-grown leaves was chemically reduced at any given irradiance. It was also observed that the quantum yield of PSII photochemistry was lower in PSI light-grown leaves. PSI light-grown leaves were more sensitive to photoinihibition and recovery was slower compared to PSII light-grown leaves, showing that the PSII reaction centre in PSI light-grown leaves was more easily impaired by photoinhibition. The increase in susceptibility of leaves to photoinhibition following blockage of chloroplast-encoded protein synthesis was greater in PSII light-grown leaves, showing that these leaves normally have a greater capacity for PSII repair. Inhibition of zeaxanthin formation by dithiothreitol slightly increased sensitivity to photoinhibition in both PSI and PSII light-grown leaves.     

Concerning a dual function of coupled cyclic electron transport in leaves

Coupled cyclic electron transport is assigned a role in the protection of leaves against photoinhibition in addition to its role in ATP synthesis. In leaves, as in reconstituted thylakoid systems, cyclic electron transport requires “poising,” i.e. availability of electrons at the reducing side of photosystem I (PSI) and the presence of some oxidized plastoquinone between photosystem II (PSII) and PSI. Under self-regulatory poising conditions that are established when carbon dioxide limits photosynthesis at high light intensities, and particularly when stomata are partially or fully closed as a result of water stress, coupled cyclic electron transport controls linear electron transport by helping to establish a proton gradient large enough to decrease PSII activity and electron flow to PSI. This brings electron donation by PSII, and electron consumption by available electron acceptors, into a balance in which PSI becomes more oxidized than it is during fast carbon assimilation. Avoidance of overreduction of the electron transport chain is a prerequisite for the efficient protection of the photosynthetic apparatus against photoinactivation.     

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.     

Xanthophyll Cycle and Inactivation of Photosystem Ⅱ Reaction Centers Alleviating Reducing Pressure to Photosystem Ⅰ in Morning Glory Leaves under Short-term High Irradiance

investigated through chlorophyll fluorescence parameters in morning glory (Ipomoea setosa) leaves, which were dipped into water, dithiothreitol (DTT) and lincomycin (LM), respectively. During the stress, both the xanthophyll cycle and D1 protein turnover could protect PSI from photoinhibition. In DTT leaves, non-photochemical quenching (NPQ) was inhibited greatly and the oxidation level of P700 (P700+) was the lowest one. However, the maximal photochemical efficiency of PSII (Fv/Fm) in DTT leaves was higher than that of LM leaves and was lower than that of control leaves. These results suggested that PSI was more sensitive to the loss of the xanthophyll cycle than PSII under high irradiance. In LM leaves, NPQ was partly inhibited, Fv/Fm was the lowest one among three treatments under high irradiance and P700+ was at a similar level as that of control leaves. These results implied that inactivation of PSII reaction centers could protect PSI from further photoinhibition. Additionally, the lowest of the number of active reaction centers to one inactive reaction center for a PSII cross-section (RC/CSo), maximal trapping rate in a PSII cross-section (TRo/CSo), electron transport in a PSII cross-section (ETo/CSo) and the highest of 1-qP in LM leaves further indicated that severe photoinhibition of PSII in LM leaves was mainly induced by inactivation of PSII reaction centers, which limited electrons transporting to PSI. However, relative to the LM leaves the higher level of RC/CSo, TRo/CSo, Fv/Fm and the lower level of 1-qP in DTT leaves indicated that PSI photoinhibition was mainly induced by the electron accumulation at the PSI acceptor side, which induced the decrease of P700+ under high irradiance.     

Effect of the anthocyanic epidermal layer on Photosystem II and I energy dissipation processes in Tradescantia pallida (Rose) Hunt

The effect of anthocyanic cells of the epidermal layer was investigated on photosynthetic activity of the higher plant Tradescantia pallida. To determine the possible indirect role of anthocyanin in photosynthesis, analysis was done on intact leaves and leaves where anthocyanic epidermal layer was removed. Energy dissipation processes related to Photosystem II (PSII) and Photosystem I (PSI) activity was done using simultaneously Chlorophyll a (Chl a) fluorescence and P700 transmittance signals change. In anthocyanic epidermal-less leaves, PSII photochemical activity was more decreased in dependence to increasing light irradiance exposure. We found that photoinhibition of PSII decreased PSI activity by reducing the electron flow toward PSI, especially under high light intensities. Under those conditions, it resulted in the accumulation of oxidized PSI reaction centers, which was stronger in leaves where the anthocyanic epidermal layer was removed. In conclusion, our results showed that the anthocyanic epidermal layer had a photoprotective effect only on the PSII and not on the PSI of T. pallida leaves, supporting the role of anthocyanin pigments in the regulation of photosynthesis for excess absorbed light irradiance.     

Photosystem II photoinhibition-repair cycle protects Photosystem I from irreversible damage

Photodamage of Photosystem II (PSII) has been considered as an unavoidable and harmful reaction that decreases plant productivity. PSII, however, has an efficient and dynamically regulated repair machinery, and the PSII activity becomes inhibited only when the rate of damage exceeds the rate of repair. The speed of repair is strictly regulated according to the energetic state in the chloroplast. In contrast to PSII, Photosystem I (PSI) is very rarely damaged, but when occurring, the damage is practically irreversible. While PSII damage is linearly dependent on light intensity, PSI gets damaged only when electron flow from PSII exceeds the capacity of PSI electron acceptors to cope with the electrons. When electron flow to PSI is limited, for example in the presence of DCMU, PSI is extremely tolerant against light stress. Proton gradient (ΔpH)-dependent slow-down of electron transfer from PSII to PSI, involving the PGR5 protein and the Cyt b6f complex, protects PSI from excess electrons upon sudden increase in light intensity. Here we provide evidence that in addition to the ΔpH-dependent control of electron transfer, the controlled photoinhibition of PSII is also able to protect PSI from permanent photodamage. We propose that regulation of PSII photoinhibition is the ultimate regulator of the photosynthetic electron transfer chain and provides a photoprotection mechanism against formation of reactive oxygen species and photodamage in PSI.