高等植物光合作用的光抑制研究进展

光抑制是目前高等植物光合作用研究中的热点,近些年来无论是对其本质的认识,还是机理研究都已取得很大进展。本文首先简要回顾了光抑制研究发展的历程,阐明现代光抑制理论包括耗散过剩光能的光保持机制运转和过剩光能对光合机构的破坏两个方面。然后,就叶黄素循环、Mehler反应、光呼吸、LHCII磷酸化、PSII光化学活性下降以及由类胡罗卜素、Cytb 559参与的一些主要光保护机制作了综述,着重论述了其作用机理及研究进展。最后,就现阶段光破坏原初作用位点的认识及光破坏机理的最新研究成果作了总结。     

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。     

光合作用的光抑制

虽然光是植物光合作用的根本推动力,没有光植物便不能进行光合作用,光不足则不能高速地进行光合作用,可是光过剩对植物来说也不是好事。当叶片接受的光能超过它所能利用的量时,光可以引起光合活性的降低。这就是光合作用的光抑制现象。它的最明显特征是光合效率的降低。在没有其它环境胁迫的条件下,晴天中午许多植物冠层表面的叶片和静止的水体表层的藻类经常发生光抑制。由于发生光抑制的前提是光能过剩,所以,任何妨碍光合作用正常进行而引起光能过剩的因素,如低温、干旱等,都会使植物易于发生光抑制。因此,在两种或两种以上环境…     

高等植物PSⅡ的光抑制与光破坏研究进展

摘要：强光对高等植物的光合作用具有抑制和破坏作用,光抑制的原初部位及主要部位在PSⅡ。综述了高等植物PSⅡ的光破坏的分子机理及其保护机制研究进展,提出了今后进一步研究的方向。     

叶黄素循环及其在光保护中的分子机理研究

植物的生命活动离不开充足的光照 ,但是当光照过强时 ,叶片吸收的光能超过了光合电子传递所需 ,过剩的光能便会对光合器官产生潜在的危害 ,引起光合作用的光抑制或光破坏。依赖于叶黄素循环的热耗散被认为是光保护的主要途径。本文着重介绍近年来有关植物叶黄素循环在酶学方面的分子调控、它的主要功能以及依赖于叶黄素循环的热耗散在光保护中的分子机理等 ,并对需进一步研究的问题作了探讨     

叶黄素循环及其在光保护中的分子机理研究

植物的生命活动离不开充足的光照,但是当光照过强时,叶片吸收的光能超过了光合电子传递所需,过剩的光能便会对光合器官产生潜在的危害,引起光合作用的光抑制或光破坏.依赖于叶黄素循环的热耗散被认为是光保护的主要途径.本文着重介绍近年来有关植物叶黄素循环在酶学方面的分子调控、它的主要功能以及依赖于叶黄素循环的热耗散在光保护中的分子机理等,并对需进一步研究的问题作了探讨.     

晴天条件下光、温变化对苹果绿色果皮原初光化学反应的影响

利用快速叶绿素荧光诱导动力学和光谱反射测定技术,研究了晴天条件下,光、温变化对苹果绿色果皮原初光化学反应的影响.结果表明：一天内,随着光、温的增强,金冠苹果果皮在12:00—14:00存在较严重的光抑制.O-J-I-P荧光诱导曲线在300 μs处的相对可变荧光（W_k）几乎没有变化,说明果皮PSⅡ的放氧复合体（OEC）的活性在一天当中没有受到强光和高温的伤害;但是果皮捕获的激子将电子传递到电子传递链中Q_A-下游电子受体的概率（Ψ_o）从8:00—12:00逐渐下降,说明金冠果皮PSⅡ反应中心受体侧的功能受到抑制.强光降低了果皮单位面积上有活性的反应中心（RC/CS）的数量,导致单位反应中心吸收的光能（ABS/RC）增加.果皮的光化学反应（TR_o/RC）不能完全利用所吸收的光能,使单位反应中心的热耗散（DI_o/RC）增加.伴随着光抑制的出现,苹果果皮叶黄素库的脱环化比例增加,表明强光下,果皮启动了叶黄素循环机制,来耗散过剩光能,以减轻过剩光能对光合机构的进一步伤害;一天中,光强和温度的增加均可加重果皮的光抑制程度,但光强对果皮的影响程度显著大于温度对果皮的影响.     

Na2CO3胁迫下星星草幼苗叶片PSⅡ光能利用和耗散与培养基质渗透势的关系

用荧光动力学的方法研究了碱性盐Na2CO3胁迫下星星草幼苗叶片PSⅡ光能利用和耗散与培养基质渗透势的关系。结果发现在大于-4bar的胁迫下,PSⅡ最大光化学效率（Fv/Fm）、PSⅡ潜在光化学效率（Fv/Fo）、PSⅡ实际光化学效率（φPSⅡ）以及开放的PSⅡ反应中心有效光化学效率（Fv’/Fm’）的变化不大;然而在小于-4bar Na2CO3胁迫下,Fv/Fm、Fv/Fo和Fv’/Fm’均随着渗透势的增大而增大,而中PSⅡ、电子传递速率（ETR）、光化学速率、捕光色素吸收的光能被用于热耗散的相对份额及热耗散速率则随着渗透势的增大而减小。这些研究结果说明星星草幼苗在Na2CO3胁迫所导致的不同的渗透胁迫下（小于-4bar和大于-4bar）其过剩光能的耗散机制可能不同,大于-4bar的胁迫下可能存在精细的渗透调节机制,而在高强度的Na2CO3所导致的渗透胁迫下具有与其它植物不同的保护机制,可能通过两条途径耗散过剩的光能,一方面通过增加捕光色素吸收的光能被用于热耗散的相对份额及热耗散速率;另一方面通过增大西PSⅡ、光化学速率、ETR,增强假循环式光合磷酸化过程,而由此引起的活性氧的增加则通过体内较高活性的保护酶系统来清除,以保护光合器官免受过剩光能的损伤。     

紫茎泽兰光合特性对生长环境光强的适应

测定了不同光强下生长的紫茎泽兰叶片最大净光合速率(Pmax)、叶绿素荧光参数、光合色素含量和比叶重(SLW),探讨了其光适应能力及生理生态学机制．强光下(100％相对光强)紫茎泽兰发生了轻度光抑制,Pmax、SLW、类胡萝卜素含量和日间热耗散升高,但热耗散能力没有提高．强光下紫茎泽兰通过：1)加强日间热耗散和活性氧清除能力以及光系统Ⅱ反应中心可逆失活来耗散过剩光能;2)增大P～以增加光能利用;3)提高SLW,降低单位干重叶绿素含量以减少光能吸收3个途径避免了光合机构光破坏．弱光下(36％、12．5％和4．5％相对光强)紫茎泽兰日间热耗散很小,SLW降低,但P～较高,这有利于其增加光能吸收和利用效率．紫茎泽兰能在很大的光强范围内有效地维持光合系统正常运转,这可能是其表现较强入侵性的原因之一．     

光合作用光抑制的研究进展

概述了植物光合作用光抑制的研究进展,包括造成光抑制和光氧化的活性氧的产生和作用机理,光抑制的作用部位,以及光保护机制等,着重从三个方面讨论了植物抗光抑制的保护机理：与光系统Ⅱ天线以及叶黄素循环相关的热耗散途径,包括光呼吸、H2O-H2O循环和环式电子传递在内的电子传递途径,以及活性氧清除机制等。     

The regulation of P700 is an important photoprotective mechanism to NaCl‐salinity in Jatropha curcas

Salinity commonly affects photosynthesis and crop production worldwide. Salt stress disrupts the fine balance between photosynthetic electron transport and the Calvin cycle reactions, leading to over‐reduction and excess energy within the thylakoids. The excess energy triggers reactive oxygen species (ROS) overproduction that causes photoinhibition in both photosystems (PS) I and II. However, the role of PSI photoinhibition and its physiological mechanisms for photoprotection have not yet been fully elucidated. In the present study, we analyzed the effects of 15 consecutive days of 100 mM NaCl in Jatropha curcas plants, primarily focusing on the photosynthetic electron flow at PSI level. We found that J. curcas plants have important photoprotective mechanisms to cope with the harmful effects of salinity. We show that maintaining P700 in an oxidized state is an important photoprotector mechanism, avoiding ROS burst in J. curcas exposed to salinity. In addition, upon photoinhibition of PSI, the highly reduced electron transport chain triggers a significant increase in H₂O₂ content which can lead to the production of hydroxyl radical by Mehler reactions in chloroplast, thereby increasing PSI photoinhibition.     

Contribution of photosynthetic electron transport,heat dissipation,and recovery of photoinactivated photosystem II to photoprotection at different temperatures in Chenopodium album leaves

Temperature dependence of photoinhibition and photoprotective mechanisms (10-35 degrees C) was investigated for Chenopodium album leaves grown at 25 degrees C under 500 micro mol quanta m(-2) s(-1). The fraction of active photosystem II (PSII) was determined after photoinhibitory treatment at different temperatures in the presence and absence of lincomycin, an inhibitor of chloroplast-encoded protein synthesis. In the absence of lincomycin, leaves were more tolerant to photoinhibition at high (25-35 degrees C) than at low (11-15 degrees C) temperatures. In the presence of lincomycin, the variation in the tolerance to photoinactivation became relatively small. The rate constant of photoinactivation (k(pi)) was stable at 25-35 degrees C and increased by 50% with temperature decrease from 25 to 11 degrees C. The rate constant of recovery of inactivated PSII (k(rec)) was more sensitive to temperature; it was very low at 11 degrees C and increased by an order of magnitude at 35 degrees C. We conclude that the recovery of photoinactivated PSII plays an essential role in photoprotection at 11-35 degrees C. Partitioning of light energy to various photoprotective mechanisms was further analyzed to reveal the factor responsible for k(pi). The fraction of energy utilized in photochemistry was lower at lower temperatures. Although the fraction of heat dissipation increased with decreasing temperatures, the excess energy that is neither utilized by photochemistry nor dissipated by heat dissipation was found to be greater at lower temperatures. The k(pi) value was strongly correlated with the excess energy, suggesting that the excess energy determines the rate of photoinactivation.     

Photoinhibition of Photosynthesis: Role of Carotenoids in Photoprotection of Chloroplast Constituents

Exposure of plants to irradiation, in excess to saturate photosynthesis, leads to reduction in photosynthetic capacity without any change in bulk pigment content. This effect is known as photoinhibition. Photoinhibition is followed by destruction of carotenoids (Cars), bleaching of chlorophylls (Chls), and increased lipid peroxidation due to formation of reactive oxygen species if the excess irradiance exposure continues. Photoinhibition of photosystem 2 (PS2) in vivo is often a photoprotective strategy rather than a damaging process. For sustainable maintenance of chloroplast function under high irradiance, the plants develop various photoprotective strategies. Cars perform essential photoprotective roles in chloroplasts by quenching the triplet Chl and scavenging singlet oxygen and other reactive oxygen species. Recently photoprotective role of xanthophylls (zeaxanthin) for dissipation of excess excitation energy under irradiance stress has been emphasised. The inter-conversion of violaxanthin (Vx) into zeaxanthin (Zx) in the light-harvesting complexes (LHC) serves to regulate photon harvesting and subsequent energy dissipation. De-epoxidation of Vx to Zx leads to changes in structure and properties of these xanthophylls which brings about significant structural changes in the LHC complex. This ultimately results in (1) direct quenching of Chl fluorescence by singlet-singlet energy transfer from Chl to Zx, (2) trans-thylakoid membrane mediated, pH-dependent indirect quenching of Chl fluorescence. Apart from these, other processes such as early light-inducible proteins, D1 turnover, and several enzymatic defence mechanisms, operate in the chloroplasts, either for tolerance or to neutralise the harmful effect of high irradiance.     

High Contents of Anthocyanins in Young Leaves are Correlated with Low Pools of Xanthophyll Cycle Components and Low Risk of Photoinhibition

We checked the hypothesis that the transient presence of anthocyanins in young leaves serves a photoprotective function. For this purpose, Rosa sp. and Ricinus communis L., whose young leaves are red to become green upon maturation, were used. Thus, young leaves with high and mature leaves with low anthocyanin contents were analysed concerning their carotenoid (Car) composition and susceptibility to photoinhibition. Cars, including the components of the xanthophyll cycle, had similar contents in young and mature leaves, when expressed on a chlorophyll basis. Yet, when expressed on a leaf area basis or on the assumed photon absorptive capacity of leaves, Cars contents were considerably lower in anthocyanic young leaves. Although this may indicate a low photodissipative potential, red young leaves were considerably less susceptible to photoinhibitory damage. The results are compatible with a photoprotective function of anthocyanins, indicating also that their presence may compensate for a low capacity in the xanthophyll cycle-dependent harmless dissipation of excess excitation energy.     

Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion.

A conserved regulatory mechanism protects plants against the potentially damaging effects of excessive light. Nearly all photosynthetic eukaryotes are able to dissipate excess absorbed light energy in a process that involves xanthophyll pigments. To dissect the role of xanthophylls in photoprotective energy dissipation in vivo, we isolated Arabidopsis xanthophyll cycle mutants by screening for altered nonphotochemical quenching of chlorophyll fluorescence. The npq1 mutants are unable to convert violaxanthin to zeaxanthin in excessive light, whereas the npq2 mutants accumulate zeaxanthin constitutively. The npq2 mutants are new alleles of aba1, the zeaxanthin epoxidase gene. The high levels of zeaxanthin in npq2 affected the kinetics of induction and relaxation but not the extent of nonphotochemical quenching. Genetic mapping, DNA sequencing, and complementation of npq1 demonstrated that this mutation affects the structural gene encoding violaxanthin deepoxidase. The npq1 mutant exhibited greatly reduced nonphotochemical quenching, demonstrating that violaxanthin deepoxidation is required for the bulk of rapidly reversible nonphotochemical quenching in Arabidopsis. Altered regulation of photosynthetic energy conversion in npq1 was associated with increased sensitivity to photoinhibition. These results, in conjunction with the analysis of npq mutants of Chlamydomonas, suggest that the role of the xanthophyll cycle in nonphotochemical quenching has been conserved, although different photosynthetic eukaryotes rely on the xanthophyll cycle to different extents for the dissipation of excess absorbed light energy.     

A major light-harvesting polypeptide of photosystem II functions in thermal dissipation

Under high-light conditions, photoprotective mechanisms minimize the damaging effects of excess light. A primary photoprotective mechanism is thermal dissipation of excess excitation energy within the light-harvesting complex of photosystem II (LHCII). Although roles for both carotenoids and specific polypeptides in thermal dissipation have been reported, neither the site nor the mechanism of this process has been defined precisely. Here, we describe the physiological and molecular characteristics of the Chlamydomonas reinhardtii npq5 mutant, a strain that exhibits little thermal dissipation. This strain is normal for state transition, high light-induced violaxanthin deepoxidation, and low light growth, but it is more sensitive to photoinhibition than the wild type. Furthermore, both pigment data and measurements of photosynthesis suggest that the photosystem II antenna in the npq5 mutant has one-third fewer light-harvesting trimers than do wild-type cells. The npq5 mutant is null for a gene designated Lhcbm1, which encodes a light-harvesting polypeptide present in the trimers of the photosystem II antennae. Based on sequence data, the Lhcbm1 gene is 1 of 10 genes that encode the major LHCII polypeptides in Chlamydomonas. Amino acid alignments demonstrate that these predicted polypeptides display a high degree of sequence identity but maintain specific differences in their N-terminal regions. Both physiological and molecular characterization of the npq5 mutant suggest that most thermal dissipation within LHCII of Chlamydomonas is dependent on the peripherally associated trimeric LHC polypeptides.     

The Effects of Excess Irradiance on Photosynthesis in the Marine Diatom Phaeodactylum tricornutum

The response of Phaeodactylum tricornutum to excess light was remarkably similar to that observed in higher plants and green algae and was characterized by complex changes in minimal fluorescence yields of fully dark-adapted samples and declines in maximum variable fluorescence levels and oxygen evolution rates. In our study the parallel decreases in the effective rate constant for photosystem II (PSII) photochemistry, the variable fluorescence yield of a dark-adapted sample, and light-limited O2 evolution rates after short (0-10 min) exposures to photoinhibitory conditions could not be attributed to damage or down-regulation of PSII reaction centers. Instead, these changes were consistent with the presence of nonphotochemical quenching of PSII excitation energy in the antennae. This quenching was analogous to that component of nonphotochemical quenching studied in higher plants that is associated with photoinhibition of photosynthesis and/or processes protecting against photoinhibition in that it did not relax readily in the dark and persisted in the absence of a bulk transthylakoid proton gradient. The quenching was most likely associated with photoprotective processes in the PSII antenna that reduced the extent of photoinhibitory damage, particularly after longer exposures. Our results suggest that a large population of damaged, slowly recovering PSII centers did not form in Phaeodactylum even after 60 min of exposure to excess actinic light.     

Photoprotective function of betacyanin in leaves of <Emphasis Type="Italic">Amaranthus cruentus</Emphasis> L. under water stress

The photoprotective function of leaf betacyanin in water-stressed Amaranthus cruentus plants was examined by comparing leaves of two strains which differ significantly in the amount of betacyanin. At 0, 1, and 2 days after the imposed water stress, leaves were subjected to high-light (HL) treatment to assess their photosynthetic capacity and photoinhibition susceptibility. The water stress equally reduced leaf relative water content (RWC), gas-exchange rate and chlorophyll (Chl) contents in both leaves, indicating that the severity of water stress was comparable between the strains. Consequently, the extent of photoinhibition after the HL treatment increased in both strains as water stress developed; however, it was significantly greater in acyanic leaves than in betacyanic leaves, suggesting lower photoinhibition susceptibility in the betacyanic strain. The betacyanic leaves also exhibited approximately 30% higher values for photochemical quenching coefficient (q_P) during the period of water stress despite the nonphotochemical quenching coefficient (q_N) did not differ significantly between the strains. These results may be partially explained by the increased amount of leaf betacyanin under water stress. Moreover, a decrease in Chl content in betacyanic leaves might have enhanced light screening effect of betacyanin by increasing relative abundance of betacyanin to Chl molecule. In addition, reduced Chl content increased light penetrability of leaves. As a result, the extent of photoinhibition at the deeper tissue was exacerbated and the Chl fluorescence emitted from these tissues was more readily detected, facilitating assessment of photoinhibition at deeper tissues where the effect of betacyanic light screening is considered to be most apparent. Our results demonstrated that leaf betacyanin contributes to total photoprotective capacity of A. cruentus leaves by lowering excitation pressure on photosystem II (PSII) via attenuation of potentially harmful excess incident light under water stress.