植物叶黄素循环与非辐射能量耗散

简述了叶黄素循环机制以及非辐射能量耗散的检测方法,并介绍了叶黄素循环与非辐射能量耗散关系的研究现状和最新进展。     

光照下叶黄素循环与非辐射能量耗散的关系

离体玉米叶片在光照下叶黄素循环与非辐射能量耗散发生明显变化.随着光强提高,玉米黄质(Z)含量显著上升,单环氧玉米黄质(A)含量在中低光强时增加但在较强光照下略有降低,紫黄质(V)含量则呈下降趋势,但叶黄素循环组分库V+A+Z上升幅度不大;相同条件下非辐射能量耗散增强,表现为非光化学猝灭荧光参数(NPQ)明显上升,同时F_v/F_m下降.分析表明,(Z+0.5A)/(V+A+Z)与NPQ呈明显正相关,而与F_v/F_m呈显著负相关,(V+0.5A)/(V+A+Z)则与之相反.由此推测,离体条件下玉米叶片中Z的环氧化和V的脱环氧化明显与非辐射能量耗散和PSⅡ光能转化过程相关.     

外源药剂处理对玉米叶黄素循环与非辐射能量耗散的影响

利用ＡｓＡ、ＤＴＴ和ＮＡＤＰＨ溶液处理离体玉米叶片,对其叶黄素循环和非辐射能量耗散都可产生一定的影响。２０ｍｍｏｌ／Ｌ ＡｓＡ可促进紫黄质（Ｖ）向单珏氧玉米黄质（Ａ）至玉米黄质（Ｚ）的转化,ＮＰＱ值和Ｆｖ／Ｆｍ均相应增加,但是生成的Ｚ在强光下（〉６５０μｍｏｌ ｍ＾－２ｓ＾－１）很容易达到饱和,５ｍｍｏｌ／Ｌ ＤＴＴ可明显抑制Ｚ的增加,但对Ａ的影响很小,同时玉米叶片的ＮＰＱ值和Ｆｖ／Ｆｍ均显著下降     

依赖叶黄素循环的非辐射能量耗散在防御珊瑚树叶片光抑制破坏中的作用

使用脉冲调制荧光仪观测了珊瑚树叶片光合作用的光抑制发生与恢复过程中几个主要荧光参数(初始荧光F_0,可变荧光与最大荧光之比F_v/F_M和非光化学荧光猝灭q_E及其快组分 q_(E—fast)、慢组分 q(E—slow))的变化,以探讨非光化学荧光猝灭不同组分的作用。 强光(约 1500μmol photons m~(-2) s~(-1))照射叶片使F_0、F_V/F_M和q_(E—fast)降低.q_(E—slow)和q_E增高。NH_4Cl处理使 F_V/F_M降低的幅度和q_E提高的幅度都增加。DTT处理使q_E水平和q_(E—slow)增加的幅度降低,而F_0和稳态荧光水平增加,强光下降低了的F_V/F_M在弱光下不易恢复。NaF处理对这些荧光参数都没有明显的影响。     

珊瑚树和大豆叶片叶绿素荧光的非光化学猝灭

用ＰＡＭ－２０００型荧光仪和７５４型分光光度计观测了珊瑚树和大豆叶片叶绿素荧光的非光化学猝灭性,中和慢３个组分和５０５ｎｍ光吸收的日变化。主要结果如下：（１）中午,珊瑚树叶片的ｑＮ８比ｑＮｆ大得多,而大豆叶片的这两个参数却几乎处于同一水平。它们的ｑＮｍ虽然也随光强变化,但与ｑＮ８和ｑＮｍ相比,除早期和傍晚以外全天的水平都是最低的。     

植物叶片和冠层光化学反射指数与叶黄素循环的关系

以一串红(Salvia splendens Ker-Gawl.)和白车轴草(Trifolium repens Linn.)为材料,使用光谱反射技术测定了这两种植物叶片水平和冠层水平的光化学反射指数(PRI)的日变化,同时使用高效液相色谱分析法测定了这两种植物叶黄素循环的日变化,分析了单叶、群体冠层的PRI与叶黄素循环和叶片实际光化学效率(ΦPSⅡ)和非光化学淬灭(NPQ)之间的关系.结果表明,不论是叶片水平还是冠层水平,两种植物PRI的变化均与叶黄素的脱环氧化程度和NPQ之间呈显著的负相关、与ΦPSⅡ呈显著的正相关.研究结果表明无论叶片水平还是冠层水平上的光谱反射指数均能非常好地反映植物光合机构对光能的利用效率.     

缺铁使大豆叶片激发能的耗散增加

缺铁叶片的光合速率大幅度下降。这种降低可能不是色素含量降低的结果 ;而且缺铁对PSII复合物的活性影响很小 ;较高的PQ还原程度显示缺铁叶片PSII受体侧电子传递受阻 ,这可能是导致光合速率下降的主要因素。强光下缺铁叶片的天线转化效率比正常叶片低 ,用于光化学反应的激发能很少。缺铁导致大豆叶片激发能耗散增加。通过抑制剂处理和叶黄素组分的分析 ,可以认为在耗散过剩激发能的过程中 ,缺铁叶片充分启动了叶黄素循环     

拟南芥叶黄素缺失突变体叶绿素荧光猝灭的特性

研究了3个叶黄素组分缺失的拟南芥核基因突变体,npq₁（缺乏玉米黄质Z和单环氧玉米黄质A）、lut₂（缺乏lutein）和lut₂-npq₁（双突变体,同时缺Z和lutein）,及其对照野生型（WT）在强光诱导下叶绿素荧光猝灭的特性.与WT相比,3个突变体的叶绿素a/b没有明显的差异,F_v/F_m则有不同幅度的增加,缺乏lutein的突变体lut₂和lut₂-npq₁的叶黄素循环库（V+A+Z）显著增大.缺乏Z的突变体npq₁和lut₂-npq₁在强光下,荧光的非光化学猝灭（NPQ）的诱导受到明显抑制,lut₂的NPQ形成也受到部分抑制.强光处理9 min后,3个突变体和WT的NPQ大小顺序为WT＞lut₂＞npq₁＞npq₁-lut₂.强光诱导过程中突变体的光化学猝灭(qP)都小于WT.强光下突变体显示较弱的抗光抑制能力,其抗光抑制能力的强弱顺序为：WT＞lut₂＞npq₁＞lut₂-npq₁.结果表明叶黄素循环不但与NPQ的形成直接相关也与qP有关.     

珊瑚树叶片叶绿素荧光非光化学猝灭的日变化和季节变化

用脉冲调制荧光仪观测了珊瑚树叶片叶绿素荧光非光化学猝灭（qE）的日变化和季节变化后发现：在晴天,qE及其慢弛豫组分（qE-slow）随着光强的增加而升高,中午达最高值,之后随光强的减弱而下降;阴天时,这两个指标的日变化不明显。在不同季节,相同日时间和同一光照强度下测定珊瑚树叶片的qE和qE-slow,两个指标在冬季明显高于春、秋两季;在短时间（1d）内改变强光下的叶片周围的温度,叶片的qE和qE-slow在高温和低温下均高于过温下测定的结果。     

植物组织中叶黄素循环组分的高效液相色谱分析法

AnalysisoftheXanthophyllCycleComponentsinPlantTissuesbyHighPer－formanceLiquidChromatographyZHAOShi－Jie,MENGQing－Wei,XUChang－Cheng,HANHong－Yan,ZOUQi（LaboratoryofPlantPhysiologyandBio-chemistry,ShandongAgriculturalUninersity,Taian271018）##D近几年来,叶黄素循环在耗散过剩光能,保护光合机构兔受强光破坏中的作用受到了普遍的关注[2,4]。在此循环中,其关键组分玉米黄质（zeaxanthin）是由紫黄质（violaxanthin）通过环氧玉米黄质（antheraxanthin）形成的。这3个组分可随光照条件的改变而…     

Non-Radiative Dissipation of Absorbed Excitation Energy Within Photosynthetic Apparatus of Higher Plants

The review deals with thermal dissipation of absorbed excitation energy within pigment-protein complexes of thylakoid membranes in higher plants. We focus on the de-excitation regulatory processes within photosystem 2 (PS2) that can be monitored as non-photochemical quenching of chlorophyll (Chl) a fluorescence consisting of three components known as energy-dependent quenching (q_E), state-transition quenching (q_T), and photoinhibitory quenching (q_I). We summarize the role of thylakoid lumen pH, xanthophylls, and PS2 proteins in q_E mechanism. Further, both the similarity between q_E and q_I and specific features of q_I are described. The other routes of thermal energy dissipation are also mentioned, that is dissipation within photosystem 1 and dissipation through the triplet Chl pathway. The significance of the individual de-excitation processes in protection against photo-oxidative damage to the photosynthetic apparatus under excess photon supply is stretched.     

Survey of Thermal Energy Dissipation and Pigment Composition in Sun and Shade Leaves

A survey was conducted of the magnitude of energy dissipationin photosystem II (expressed as nonphotochemi-cal quenchingof chlorophyll fluorescence, NPQ) as well as leaf carotenoidcomposition of a wide range of different plant species growingin deep shade and/or full sun. Consistently higher levels ofthe reversible component of NPQ as well as higher degrees ofrapidly attainable de-epoxida-tion of the xanthophyll cycle(VAZ) pool were observed in sun leaves compared to deep shadeleaves. It is concluded that these altered features of the xanthophyllcycle allowed sun leaves to dissipate excess energy more effectivelyover the short term. In addition to the rapid increase in reversibleNPQ, shade leaves exhibited a slow further, and sustained, increasein NPQ. In contrast to these deep shade leaves experimentallyexposed to high PFDs, understory leaves experiencing highlyvariable PFD in their natural environment appeared to be ableto dissipate excess excitation energy adequately via xanthophyllcycle-dependent thermal dissipation. Furthermore, very consistenttrends across plant species were observed for changes in carotenoidcomposition (pools of carotenes, VAZ, and other xantho-phylls)in response to light environment, as long as it is assumed thatin some species rß-carotene can be replaced by     

Diversity in Xanthophyll Cycle Pigments Content and Related Nonphotochemical Quenching (NPQ) Among Microalgae: Implications for Growth Strategy and Ecology

Xanthophyll cycle-related nonphotochemical quenching (NPQ), which is present in most photoautotrophs, allows dissipation of excess light energy. Xanthophyll cycle-related NPQ depends principally on xanthophyll cycle pigments composition and their effective involvement in NPQ. Xanthophyll cycle-related NPQ is tightly controlled by environmental conditions in a species-/strain-specific manner. These features are especially relevant in microalgae living in a complex and highly variable environment. The goal of this study was to perform a comparative assessment of NPQ ecophysiologies across microalgal taxa in order to underline the specific involvement of NPQ in growth adaptations and strategies. We used both published results and data acquired in our laboratory to understand the relationships between growth conditions (irradiance, temperature, and nutrient availability), xanthophyll cycle composition, and xanthophyll cycle pigments quenching efficiency in microalgae from various taxa. We found that in diadinoxanthin-containing species, the xanthophyll cycle pigment pool is controlled by energy pressure in all species. At any given energy pressure, however, the diatoxanthin content is higher in diatoms than in other diadinoxanthin-containing species. XC pigments quenching efficiency is species-specific and decreases with acclimation to higher irradiances. We found a clear link between the natural light environment of species/ecotypes and quenching efficiency amplitude. The presence of diatoxanthin or zeaxanthin at steady state in all species examined at moderate and high irradiances suggests that cells maintain a light-harvesting capacity in excess to cope with potential decrease in light intensity.     

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.     

依赖于叶黄素循环的热耗散对茶树叶片防御强光破坏的相关试验研究

经叶黄素循环抑制剂——二硫苏糖醇(DIT)处理的茶树叶片,以850μmol．m^-2．s^-1的PFD照射120min后,福鼎大白茶的叶黄素循环组分中的环氧玉米黄素(A)和玉米黄素(Z)含量之和降低了76．5％,结果导致非光化学猝灭(NPQ)、光系统Ⅱ(PSⅡ)的光化学效率(Fv／Fm)、光化学猝灭系数(qP)、PSⅡ实际光化学量子效率(ψPSⅡR)和光合电子传递速率(ETR)明显下降,而F0显著上升,暗恢复后Fv／Fm恢复程度小于未经DIT处理的叶片。自然光强下,NPQ与与叶黄素循环的脱环氧化程度(A Z)/(V A Z)比值呈明显的正线性关系(R=0．9488^***)。这些结果充分证明依赖与叶黄素循环的热耗散是茶树叶片光合器官防御强光破坏的主要途径。     

Enhanced Employment of the Xanthophyll Cycle and Thermal Energy Dissipation in Spinach Exposed to High Light and N Stress

The involvement of the xanthophyll cycle in photoprotection of N-deficient spinach (Spinacia oleracea L. cv Nobel) was investigated. Spinach plants were fertilized with 14 mM nitrate (control, high N) versus 0.5 mM (low N) fertilizer, and grown under both high- and low-light conditions. Plants were characterized from measurements of photosynthetic oxygen exchange and chlorophyll fluorescence, as well as carotenoid and cholorophyll analysis. Compared with the high-N plants, the low-N plants showed a lower capacity for photosynthesis and a lower chlorophyll content, as well as a lower rate of photosystem II photosynthetic electron transport and a corresponding increase in thermal energy dissipation activity measured as nonphotochemical fluorescence quenching. The low-N plants displayed a greater fraction of the total xanthophyll cycle pool as zeaxanthin and antheraxanthin at midday, and an increase in the ratio of xanthophyll cycle pigments to total chlorophyll. These results indicate that under N limitation both the light-collecting system and the photosynthetic rate decrease. However, the increased dissipation of excess energy shows that there is excess light absorbed at midday. We conclude that spinach responds to N limitation by a combination of decreased light collection and increased thermal dissipation involving the xanthophyll cycle.     

The peculiar NPQ regulation in the stramenopile Phaeomonas sp. challenges the xanthophyll cycle dogma

In changing light conditions, photosynthetic organisms develop different strategies to maintain a fine balance between light harvesting, photochemistry, and photoprotection. One of the most widespread photoprotective mechanisms consists in the dissipation of excess light energy in the form of heat in the photosystem II antenna, which participates to the Non Photochemical Quenching (NPQ) of chlorophyll fluorescence. It is tightly related to the reversible epoxidation of xanthophyll pigments, catalyzed by the two enzymes, the violaxanthin deepoxidase and the zeaxanthin epoxidase. In Phaeomonas sp. (Pinguiophyte, Stramenopiles), we show that the regulation of the heat dissipation process is different from that of the green lineage: the NPQ is strictly proportional to the amount of the xanthophyll pigment zeaxanthin and the xanthophyll cycle enzymes are differently regulated. The violaxanthin deepoxidase is already active in the dark, because of a low luminal pH, and the zeaxanthin epoxidase shows a maximal activity under moderate light conditions, being almost inactive in the dark and under high light. This light-dependency mirrors the one of NPQ: Phaeomonas sp. displays a large NPQ in the dark as well as under high light, which recovers under moderate light. Our results pinpoint zeaxanthin epoxidase activity as the prime regulator of NPQ in Phaeomonas sp. and therefore challenge the deepoxidase-regulated xanthophyll cycle dogma.     

Leaf Xanthophyll content and composition in sun and shade determined by HPLC

As a part of our investigations to test the hypothesis that zeaxanthin formed by reversible de-epoxidation of violaxanthin serves to dissipate any excessive and potentially harmful excitation energy we determined the influence of light climate on the size of the xanthophyll cycle pool (violaxanthin + antheraxanthin + zeaxanthin) in leaves of a number of species of higher plants. The maximum amount of zeaxanthin that can be formed by de-epoxidation of violaxanthin and antheraxanthin is determined by the pool size of the xanthophyll cycle. To quantitate the individual leaf carotenoids a rapid, sensitive and accurate HPLC method was developed using a non-endcapped Zorbax ODS column, giving baseline separation of lutein and zeaxanthin as well as of other carotenoids and Chl a and b.The size of the xanthophyll cycle pool, both on a basis of light-intercepting leaf area and of light-harvesting chlorophyll, was ca. four times greater in sun-grown leaves of a group of ten sun tolerant species than in shade-grown leaves in a group of nine shade tolerant species. In contrast there were no marked or consistent differences between the two groups in the content of the other major leaf xanthophylls, lutein and neoxanthin. Also, in each of four species examined the xanthophyll pool size increased with an increase in the amount of light available during leaf development whereas there was little change in the content of the other xanthophylls. However, the -carotene/-carotene ratio decreased and little or no -carotene was detected in sun-grown leaves. Among shade-grown leaves the -carotene/-carotene ratio was considerably higher in species deemed to be umbrophilic than in species deemed to be heliophilic.The percentage of the xanthophyll cycle pool present as violaxanthin (di-epoxy-zeaxanthin) at solar noon was 96–100% for shade-grown plants and 4–53% for sun-grown plants with zeaxanthin accounting for most of the balance. The percentage of zeaxanthin in leaves exposed to midday solar radiation was higher in those with low than in those with high photosynthetic capacity.The results are consistent with the hypothesis that the xanthophyll cycle is involved in the regulation of energy dissipation in the pigment bed, thereby preventing a buildup of excessive excitation energy at the reaction centers.Abbreviations A antheraxanthin - C -carotene - C -carotene - EPS epoxidation state (V+0.5A)/(V+A+Z) - L lutein - N neoxanthin - PFD photon flux density - V violaxanthin - Z zeaxanthin C.I.W.-D.P.B. Publiation No. 1035