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。     

阳生植物和阴生植物的叶黄素循环与非辐射能量耗散X

自然条件下阳生植物和阴生植物的光合速率存在着明显的差距,它们拥有不同的适应强光胁迫的能力,前者明显强于后者。从叶黄素组分来看,阳生植物拥有更大的叶黄素库[紫黄质（V）＋单环环氧玉米黄质（A）＋玉米黄质（Z）],其中Z和[Z＋A]的含量更明显高于阴生植物;从阳生植物或阴生植物内部来看,不同物种间,Z1[Z＋A]和[V＋A＋Z]含量的差异相对较小,A则基本相同;不论是阳生植物还是阴生植物,非光化学猝灭值与Z、[Z＋A]及[V＋A＋Z]含量均呈现较好的正相关关系,后三者含量越高,非光化学猝灭值越大,而且[V＋A＋Z]库的大小与Z含量基本上是成比例增另的。说明在不同植物种间,[Z＋]（主要是Z）仍然是影响非辐射能量耗能的主要因素。     

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。     

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。     

链霉素处理对玉米叶片叶绿素荧光参数和叶黄素脱环氧化水平的影响

利用叶绿素荧光分析技术和高效液相色谱研究了链霉素(SM,叶绿体基因编码蛋白的抑制剂)处理玉米叶片的叶黄素循环及依赖叶黄素循环的热耗散。与对照相比,强光下SM处理叶片的最大光化学效率(Fv／Fm)降低且不能完全恢复,同时电子传递速率(ETR)显著下降。而且,SM处理叶片的非光化学淬灭(NPQ)和叶黄素循环的脱环氧化水平增加。但是,NPQ的主要组分高能态(qE)淬灭减小。因此,推测qE的降低可能与电子传递速率降低有关。     

田间小麦叶片光合作用的光抑制不伴随D1蛋白的净降解

通过测定田间小麦（Ｔｒｉｔｉｃｕｍ ａｅｓｔｉｖｕｍ ）叶片Ｄ１蛋白的含量、光合放氧和叶绿素ａ 荧光,探讨了叶片光合作用的光抑制与Ｄ１蛋白净降解的关系。田间的小麦叶片受到晴天中午光照约３ ｈ 以后,表观光合量子效率（Φ）、光系统Ⅱ的光化学效率（Ｆｖ／Ｆｍ ）和初始荧光（Ｆ０）明显下降;若将叶片转入弱光下,这３个指标可在１ ｈ 内基本恢复;强光照射过程中Ｄ１蛋白的含量没有显著变化;Ｄ１蛋白合成抑制剂ＳＭ 使强光下叶片的慢驰豫的非光化学荧光猝灭（ｑＥ－ｓｌｏｗ ）明显增加;在弱光下恢复时引入链霉素（ＳＭ）不影响叶片光合功能的恢复;用二硫苏糖醇（ＤＴＴ）抑制叶黄素循环使中午强光照射后的叶片中Ｄ１蛋白的含量降低３０％ 左右。这些结果都表明,田间小麦叶片光合作用的光抑制不是由于Ｄ１蛋白的净降解,而是由于非辐射能量耗散的增加引起的。     

叶黄素循环酶

依赖叶黄素循环的热耗散是一种主要防御光破坏的机制。参与叶黄素循环的酶是紫黄质脱环氧化酶和玉米黄质环氧化酶 ,紫黄质脱环氧化酶已分离纯化 ,其 c DNA已被克隆 ,其活性主要受跨类囊体膜的 p H梯度和抗坏血酸浓度的调节 ;玉米黄质环氧化酶还没有被分离出来 ,但其 c DNA也已被克隆 ;其活性主要与NADPH的浓度、O2 及光等有关。     

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     

Linear models relating xanthophylls and lumen acidity to non-photochemical fluorescence quenching. Evidence that antheraxanthin explains zeaxanthin-independent quenching

Zeaxanthin has been correlated with high-energy non-photochemical fluorescence quenching but whether antheraxanthin, the intermediate in the pathway from violaxanthin to zeaxanthin, also relates to quenching is unknown. The relationships of zeaxanthin, antheraxanthin and pH to fluorescence quenching were examined in chloroplasts ofPisum sativum L. cv. Oregon andLactuca sativa L. cv. Romaine. Data matrices as five levels of violaxanthin de-epoxidation against five levels of light-induced lumen-proton concentrations were obtained for both species. The matrices included high levels of antheraxanthin as well as lumen-proton concentrations induced by subsaturating to saturation light levels. Analyses of the matrices by simple linear and multiple regression showed that quenching is predicted by models where the major independent variable is the product of lumen acidity and de-epoxidized xanthophylls, the latter as the sum of zeaxanthin and antheraxanthin. The interactions of lumen acidity and xanthophyll concentration are shown in three-dimensional plots of the best-fit multiple regression models. Antheraxanthin apparently contributes to quenching as effectively as zeaxanthin and explains quenching previously not accounted for by zeaxanthin. Hence, we propose that all high-energy dependent quenching is xanthophyll dependent. Quenching requires a threshold lumen pH that varies with xanthophyll composition. After the threshold, quenching is linear with lumen acidity or xanthophyll composition.     

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.     

Characterization of acclimation of Hordeum vulgare to high irradiation based on different responses of photosynthetic activity and pigment composition

The ability of spring barley (Hordeum vulgare cv. Akcent) to adjust the composition and function of the photosynthetic apparatus to growth irradiances of 25–1200 μmol m⁻² s⁻¹ was studied by gas exchange and chlorophyll a fluorescence measurements and high-performance liquid chromatography. The increased growth irradiance stimulated light- and CO₂-saturated rates of CO₂ assimilation expressed on a leaf area basis up to 730 μmol m⁻² s⁻¹ (HL730), whereas at an irradiance of 1200 μmol m⁻² s⁻¹ (EHL1200) both rates decreased significantly. Further, the acclimation to EHL1200 was associated with an extremely high chlorophyll a/b ratio (3.97), a more than doubled xanthophyll cycle pool (VAZ) and a six-fold higher de-epoxidation state of the xanthophyll cycle pigments as compared to barley grown under 25 μmol m⁻² s⁻¹ (LL25). EHL1200 plants also exhibited a long-term inhibition of Photosystem II (PS II) photochemical efficiency (F _v/F _m). Photosynthetic capacity, chlorophyll a/b and VAZ revealed a linear trend of dependence on PS II excitation pressure in a certain range of growth irradiances (100–730 μmol m⁻² s⁻¹). The deviation from linearity of these relationships for EHL1200 barley is discussed. In addition, the role of increased VAZ and/or accumulation of zeaxanthin and antheraxanthin in acclimation of barley to high irradiance is studied with respect to regulation of non-radiative dissipation and/or photochemical efficiency within PS II. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nocturnally retained zeaxanthin does not remain engaged in a state primed for energy dissipation during the summer in two Yucca species growing in the Mojave Desert

Differently oriented leaves of Yucca schidigera and Yucca brevifolia were characterized in the Mojave Desert with respect to photosystem II and xanthophyll cycle activity during three different seasons, including the hot and dry summer, the relatively cold winter, and the mild spring season. Photosynthetic utilization of a high percentage of the light absorbed in PSII was observed in all leaves only during the spring, whereas very high levels of photoprotective, thermal energy dissipation were employed both in the summer and the winter season in all exposed leaves of both species. Both during the summer and the winter season, when energy dissipation levels were high diurnally, xanthophyll cycle pools (relative to either Chl or other carotenoids) were higher relative to the spring, and a nocturnal retention of high levels of zeaxanthin and antheraxanthin (Z + A) occurred in all exposed leaves of both species. Although this nocturnal retention of Z + A was associated with nocturnal maintenance of a low PSII efficiency (F_v/F_m) on a cold winter night, pre‐dawn F_v/F_m was high in (Z + A)‐retaining leaves following a warm summer night. This indicates nocturnal engagement of Z + A in a state primed for energy dissipation throughout the cold winter night – while high levels of retained Z + A were not engaged for energy dissipation prior to sunrise on a warm summer morning. Possible mechanisms for a lack of sustained engagement of retained Z + A for energy dissipation at elevated temperatures are discussed.