Stimulation of the diadinoxanthin cycle by UV-B radiation in the diatom Phaeodactylum tricornutum

In this study we show that the diadinoxanthin cycle in the diatom Phaeodactylum tricornutum is stimulated by mild UV-B radiation. High steady state concentrations of diatoxanthin established during a period of strong actinic illumination with white light (300 mol photons m-2 s-1 PAR) are further increased if weak UV-B (3 mol photons m-2 s-1) is additionally applied. Short term increases in the diatoxanthin concentration caused by UV-B strongly correlate with a stoichiometric decrease in diadinoxanthin. The UV-B dependent increase in diatoxanthin is correlated with a concommitant enhancement of non-photochemical quenching of chlorophyll fluorescence and a decrease in the quantum efficiency of oxygen evolution. This indicates that UV-B induced diatoxanthin functions in thermal energy dissipation. Possible scenarios for a stimulation of the diadinoxanthin cycle by UV-B are discussed.     

Photoprotection in a zeaxanthin- and lutein-deficient double mutant of Arabidopsis

When light absorption by a plant exceeds its capacity for light utilization, photosynthetic light harvesting is rapidly downregulated by photoprotective thermal dissipation, which is measured as nonphotochemical quenching of chlorophyll fluorescence (NPQ). To address the involvement of specific xanthophyll pigments in NPQ, we have analyzed mutants affecting xanthophyll metabolism in Arabidopsis thaliana. An npq1 lut2 double mutant was constructed, which lacks both zeaxanthin and lutein due to defects in the violaxanthin de-epoxidase and lycopene -cyclase genes. The npq1 lut2 strain had normal Photosystem II efficiency and nearly wild-type concentrations of functional Photosystem II reaction centers, but the rapidly reversible component of NPQ was completely inhibited. Despite the defects in xanthophyll composition and NPQ, the npq1 lut2 mutant exhibited a remarkable ability to tolerate high light.This revised version was published online in October 2005 with corrections to the Cover Date.     

GENERAL FEATURES OF PHOTOPROTECTION BY ENERGY DISSIPATION IN PLANKTONIC DIATOMS (BACILLARIOPHYCEAE)1

Planktonic diatoms (Bacillariophyceae) have to cope with large fluctuations of light intensity and periodic exposure to high light. After a shift to high light, photoprotective dissipation of excess energy characterized by the nonphotochemical quenching of fluorescence (NPQ) and the concomitant deepoxidation of diadinoxanthin to diatoxanthin (DT) were measured in four different planktonic marine diatoms (Bacillariophyceae): Skeletonema costatum (Greville) Cleve, Cylindrotheca fusiformis Reimann et Lewin, Thalassiosira weissflogii (Grunow) Fryxell et Hasle, and Ditylum brightwellii (West) Grunow in comparison to the model organism Phaeodactylum tricornutum Böhlin. Upon a sudden increase of light intensity, deepoxidation was rapid and de novo synthesis of DT also occurred. In all species, NPQ was linearly related to the amount of DT formed during high light. In this report, we focused on the role of DT in the dissipation of energy that takes place in the light‐harvesting complex. In S. costatum for the same amount of DT, less NPQ was formed than in P. tricornutum and as a consequence the photoprotection of PSII was less efficient. The general features of photoprotection by harmless dissipation of excess energy in planktonic diatoms described here partly explain why diatoms are well adapted to light intensity fluctuations.     

植物光破坏防御机制研究进展

光破坏防御机制是植物为应对复杂多变的自然环境而产生的保护措施,这些措施从形态、生理和生化等方面反映了植物对环境的适应能力。本文根据光抑制的机理,对近年来植物的光破坏防御机制以及高等植物叶黄素循环机制的研究现状进行综述,认为叶黄素循环防御机制是植物光保护作用的重要措施之一。     

STREAM PERIPHYTON PHOTOACCLIMATION RESPONSE IN FIELD CONDITIONS: EFFECT OF COMMUNITY DEVELOPMENT AND SEASONAL CHANGES1

The photochemical behavior of intact stream periphyton communities in France was evaluated in response to the time course of natural light. Intact biofilms grown on glass substrata were collected at three development stages in July and November, and structural parameters of the biofilms were investigated (diatom density and taxonomy). At each season, physiological parameters based on pigment analysis (HPLC) and pulse‐amplitude‐modulated (PAM) chl fluorescence technique were estimated periodically during a day from dawn to zenith. Regardless of the community studied, the optimal quantum yield of PSII (F_v/F_m), the effective PSII efficiency (Φ_PSII), the nonphotochemical quenching (NPQ), and the relative electron transport rate (rETR) exhibited clear dynamic patterns over the morning. Moreover, microalgae responded to the light increase by developing the photoprotective xanthophyll cycle. The analysis of P‐I parameters and pigment profiles suggests that July communities were adapted to higher light environments in comparison with November ones, which could be partly explained by a shift in the taxonomic composition. Finally, differences between development stages were significant only in July. In particular, photoinhibition was less pronounced in mature assemblages, indicating that self‐shading (in relation to algal biomass) could have influenced photosynthesis in older communities.     

Pigments, photosynthesis and photoinhibition in two amphibious plants: consequences of varying carbon availability

In the present study, we investigated the effects of CO(2) availability on photosynthesis, photoinhibition and pigmentation in two species of amphibious plants, Lobelia cardinalis and Nesaea crassicaulis. The plants were grown emergent under atmospheric conditions and submerged under low and high CO(2) availability. Compared with Lobelia, Nesaea had thin leaves and few stomata in all CO(2) treatments. While Lobelia expressed no variation in anthocyanin content among treatments, Nesaea produced high concentrations of anthocyanin when submerged. Lobelia photosynthesis increased in response to increasing CO(2) availability, and photoinhibition was negatively related to xanthophyll content. By contrast, Nesaea photosynthesis was highest under submerged conditions, and there was no relationship between photoinhibition and the xanthophyll content. We conclude that the response of Lobelia to varying CO(2) availability is similar to that of terrestrial plants and that this species relies on the xanthophyll cycle for nonphotochemical quenching (NPQ) and protection against photoinhibition. By contrast, the thin leaves, few stomata and low levels of chlorophylls and accessory pigments in Nesaea, relative to Lobelia, suggest adaptation to a submerged habitat. While Nesaea does not seem to rely on the xanthophyll cycle or other xanthophylls for NPQ, some role of anthocyanins in the protection against photoinhibition cannot be ruled out, owing to its effect as a sunscreen and as an efficient quencher of free radicals.     

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.     

The giant kelp Macrocystis pyrifera presents a different nonphotochemical quenching control than higher plants

Here the mechanisms involved in excitation energy dissipation of Macrocystis pyrifera were characterized to explain the high nonphotochemical quenching of chlorophyll a (Chla) fluorescence (NPQ) capacity of this alga. We performed a comparative analysis of NPQ and xanthophyll cycle (XC) activity in blades collected at different depths. The responses of the blades to dithiothreitol (DTT) and to the uncoupler NH4Cl were also assayed. The degree of NPQ induction was related to the amount of zeaxanthin synthesized in high light. The inhibition of zeaxanthin synthesis with DTT blocked NPQ induction. A slow NPQ relaxation upon the addition of NH4Cl, which disrupts the transthylakoid proton gradient, was detected. The slow NPQ relaxation took place only in the presence of de-epoxidated XC pigments and was related to the epoxidation of zeaxanthin. These results indicate that in M. pyrifera, in contrast to higher plants, the transthylakoid proton gradient alone does not induce NPQ. The role of this gradient seems to be related only to the activation of the violaxanthin de-epoxidase enzyme.     

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.     

Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest

The influence of the growth photon flux density (PFD) on the size and composition of the carotenoid pool and the size of the reduced ascorbate pool was determined across a light gradient from the forest floor to the canopy and the forest edge of a sub-tropical rainforest in New South Wales, Australia. Nineteen plant species (most collected from multiple sites) representing a broad taxonomic range consistently possessed larger total carotenoid pools when found growing in more exposed sites. There was a significant positive correlation between β-carotene content and growth PFD and a significant negative correlation between α-carotene content and growth PFD. Neoxanthin content exhibited no significant trend while the trend in lutein content varied with mode of expression. The pigments of the xanthophyll cycle (violaxanthin, antheraxanthin and zeaxanthin) exhibited the most pronounced response to growth PFD; they comprised a much greater portion of the total carotenoid pool in high light-acclimated plants. The pool of reduced ascorbate was also several-fold greater in high light-acclimated plants. These acclimatory changes in carotenoid and ascorbate content are consistent with a need for a greater capacity to dissipate excessive absorbed light energy in high light-acclimated plants.     

遮荫棉花转入强光后光合作用的光抑制及其恢复

研究了遮荫棉花（Gossypium hirsutum L.）突然由遮荫条件暴露在自然强光下时,叶绿素荧光发射、叶绿体色素组成、净光合速率（Pn）等在光照转移当天以及随后的适应过程中（光照转换后15d内）的变化。遮荫棉花突然转到强光下,叶片发生了严重的光合作用光抑制,叶绿素荧光参数Fv/Fm和φPSⅡ急剧降低,且明显低于自然光照下生长的叶片,而F。值却明显升高。这些参数即使在光照转换的次日清晨也不能完全恢复。Fv/Fm和Pn在光照转换以后的4d内持续降低,在第6天以后开始逐渐升高,在10-12d达到稳定值,表现出遮荫棉花叶片对光强变化的一定适应性,但Fv/Fm和Pn均未达到自然光照条件下生长的棉花叶片的相应值。最后的Pn值较遮荫下叶片增加60%,但同自然光照下生长的叶片相比只有后的40%。试验结果还表明,光照转换以后叶片内叶黄素循环库逐渐增大,在较短的时间内（3d）即可达到较高的水平。遮荫棉花突然转么自然强光下,叶片Fv/Fm及Pn的降低与PSⅡ反应中心的破坏有关,在对强光的适应过程中依赖叶黄素循环的热耗散等保护机制增强。光保护机制的逐渐完善有助于减轻叶片由遮荫转到强光下遭受的光破坏。     

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