光和水分胁迫对臭柏实生幼苗光化学效率及色素组成的影响

为探明毛乌素沙地3年生臭柏(Sabina vulgaris)实生苗在不同光照和水分条件下的光抑制响应机制,研究了各处理臭柏实生苗的最大光化学效率(F_v/F_m)及叶绿素(Chla+Chlb)和叶黄素(A+V+Z)含量,分析了其叶绿素循环和叶黄素循环的变化规律。结果表明,77%透光区通过减少Chlb含量,升高Chla/Chlb,避免光能过剩;同时,增加A+V+Z及热散逸色素(A+Z)含量、提高(A+V+Z)/(Chla+Chlb)和(A+V)/(A+V+Z)值,耗散过剩光能,避免光破坏。25%透光区的叶绿素和叶黄素循环机制随着水分条件的变化迅速发生改变。10%透光区通过增加Chlb含量,降低Chla/Chlb,捕捉更多的光能,几乎不存在光抑制。毛乌素臭柏实生幼苗能够适应不同的光照和水分条件,在恶劣的沙漠中完成天然更新,形成独特的群落景观,与叶绿素循环和叶黄素循环有着密切的关系。     

大气中二氧化碳浓度升高对光合作用的影响(下)

ＣＯ２浓度加倍对光合色素含量的影响ＣＯ２浓度加倍有利于植物叶片单位鲜重或单位叶面积的叶绿素和类胡萝卜素含量的提高。叶绿素含量的提高,显然有助于植物捕获更多光能供光合作用所利用。因为在ＣＯ２浓度加倍条件下,植物要充分利用环境资源,增加对ＣＯ２的同化,需要通过增加叶片叶绿素的含量,或扩大叶面积来提高对光能的捕获能力,以满足碳同化时能量的需求。此外,ＣＯ２浓度加倍;能降低叶绿素ａ／ｂ比值,说明它更有利形成叶绿素ｂ。以含等量叶绿素的叶绿体所作的实验表明,来自生长在ＣＯ２浓度加倍条件下的植物叶绿体,对光能…     

两种杜鹃花属植物对长期遮阴后全光照环境的生理响应及其光保护机制

植物通过提高光能利用能力和光保护途径以响应环境光强的增加, 但不同植物对环境光强增加的生理响应存在差异, 从而导致植物对光环境的适应性不一致。为探讨植物对光环境变化的生理响应及其适应机制, 该文以遮阴条件下培养1年的2种杜鹃属(Rhododendron)植物比利时杜鹃(R. hybrida)和杜鹃(R. simsii)为材料, 对其由遮阴后转入全光照下培养5天时的叶片叶绿素荧光参数及其快速光曲线变化进行了比较研究, 以期从叶片吸收光能分配和光保护机制的角度探讨这2种植物对光环境变化的适应机制。结果表明: 全光照降低了喜阴植物比利时杜鹃叶片的光化学反应和热耗散能力, 且其吸收光能分配于光化学反应和调节性能量耗散部分的比例减少, 导致光系统II反应中心过量激发能积累, 造成了叶片光抑制甚至光破坏。杜鹃作为耐阴喜光植物对光环境变化具有较强的适应性, 具有较高的光化学反应、热耗散和环式电子传递能力等内在生理特性; 在遮阴和全光照两种光环境下均能维持较高的吸收光能在光化学反应和调节性能量耗散部分的分配比例, 从而保护了光合机构的正常运行, 是其全光照强光未造成叶片光抑制的原因。     

光强对三种喀斯特植物幼苗生长和光合特性的影响

以块根紫金牛、地枫皮和秀丽海桐的2年生幼苗为材料,通过搭建遮阴棚设置3个光照强度,对其不同光强下的形态结构、生物量分配以及光合特性进行了比较研究,以探讨光强对喀斯特植物幼苗生长和光合特性的影响。结果表明:遮阴促进了3种喀斯特植物幼苗株高、总叶面积和冠面积等形态指标的增长。总体而言,3种植物幼苗的根生物量比和根冠比均随光强的增大而显著增大,叶生物量比、叶面积比和比叶面积均随光强的增大而显著降低。强光显著降低了地枫皮与块根紫金牛的净光合速率(Pn),非气孔限制是其Pn下降的主要因素;秀丽海桐的Pn随光强的升高而显著增大。遮阴显著增加了地枫皮和秀丽海桐的单位面积叶绿素含量,强光显著增加了块根紫金牛的单位面积类胡萝卜素含量、叶绿素a/b和类胡萝卜素/叶绿素比。地枫皮和块根紫金牛在50%遮阴强度下的总生物量最大,秀丽海桐总生物量随光强增大而显著增大。3种喀斯特植物幼苗对不同的光环境表现出较强的适应性,但强光不利于地枫皮和块根紫金牛幼苗的生长,而秀丽海桐幼苗对光的适应性很强。     

光质对花生幼苗叶片光合色素含量及光合特性的影响

在单色LED灯光照条件下,以青花6号花生品种为材料,研究了不同光质对花生幼苗光合色素含量及光合特性的影响.结果表明:与自然光照相比,蓝光(445~470 nm)可显著提高花生幼苗比叶面积、叶绿素a/b值和类胡萝卜素含量,净光合速率、气孔导度、蒸腾速率较高,胞间CO2浓度较低,光合效率显著提高;红光(610~660 nm)显著提高了叶片叶绿素含量,降低了比叶面积、叶绿素a/b值和类胡萝卜素含量,光合效率低于自然光照;绿光(515~520 nm)和黄光(590~595 nm)不利于光合色素的积累,显著抑制了花生幼苗叶片的光合作用.     

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     

遮荫对紫叶李幼苗叶片色素含量及光合速率的影响

以露地栽培的1年生紫叶李和绿叶李为试材,研究了全光照(CK)、中度遮荫(透光率为43%)、重度遮荫(透光率为27%)处理下叶片色素含量及净光合速率的变化.结果表明:在夏季,各处理紫叶李幼苗叶片叶绿素含量、类胡萝卜素含量随遮荫时间的延长逐渐升高,花色素苷含量、花色素苷含量/叶绿素含量比值降低;在秋季,各处理紫叶李幼苗叶片叶绿素含量、类胡萝卜素含量先升高后降低,而花色素苷含量、花色素苷含量/叶绿素含量比值则先降低后升高.在夏秋两季,紫叶李3种色素含量均高于绿叶李,全光照下的紫叶李叶片花色素苷含量、花色素苷含量/叶绿素含量比值及净光合速率日积分值均高于遮荫处理.紫叶李具备一定的喜光性,全光照环境有利于其彩色的呈现和光合性能的发挥.     

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.     

测定叶绿素的几种方法

在植物界,从属于原核生物的蓝藻到高等植物,都具有叶绿素,只不过它们所含的叶绿素的量和种类不同而已。叶绿素共有 a、b、c 和 d四种,在绿色植物中主要是叶绿素 a 和 b 两种。它们(还有类胡萝卜素)在植物体内与蛋白质构成色素蛋白复合体,负责对光能的吸收和传递,并最后把吸收的光能汇集到光合作用的“反应中心”(reaction center)。人们通常把前者称为天线色素。“反应中心”是处于特殊状态的     

两个不同基因型小麦光抑制特性的比较

比较了两个不同基因型小麦(Triticum aestivum L.)"京411"和"小偃54"的原初光能转化效率、荧光猝灭参数和光合色素对强光胁迫的响应.在正常生长条件下"京411"的光合色素含量高于"小偃54";但在高光强下"京411"出现明显的光抑制,而"小偃54"对高光强的适应上优于"京411"."小偃54"适应高光强的原因是它在高光强下能大幅度地提高叶黄素循环的调控因子抗坏血酸的浓度及紫黄素脱环氧化酶(vDE)的活性,从而加速叶黄素循环对过多光能的耗散过程.     

Relationships between carotenoid composition and growth habit in British plant species

The pigment composition of leaves from a number of different plant species collected from field sites in the region of Sheffield, UK, have been compared using high-performance liquid chromatography. Expression of pigment content per unit leaf area was dominated by variation in the total leaf chlorophyll. Neither chlorophyll per unit area nor the chlorophyll a/b ratio were found to be correlated with the habitat from which the plants originated. When the amounts of different carotenoids were expressed relative to the total carotenoid pool, it was found that whilst neither total carotene (α- +β-carotene) nor neoxanthin correlated with ability to grow in shade, the leaf content of both lutein and the total xanthophyll cycle carotenoids (zeaxanthin, anther-axanthin and violaxanthin) did, with lutein content being high in shade species and xanthophyll cycle intermediates low. There was a strong negative correlation between the relative amounts of each of these groups of carotenoids. The ratio of lutein to xanthophyll cycle carotenoids was strongly correlated to an index of shade tolerance.     

Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis

The photosynthetic pigments of twigs in five tree and shrub species possessing chlorenchyma under a well developed, stomata-less, and highly photon absorptive periderm were analysed and compared to those of the corresponding canopy leaves. We asked whether the unavoidable shade acclimation of corticular chlorenchyma results in photosynthetic pigment complements typically found in shade leaves. As expected, chlorophyll (Chl) a/b ratios in twigs were consistently low. However, carotenoid (Car) analysis did not confirm the initial hypothesis, since twigs generally contained increased Chl-based pool sizes of the xanthophyll cycle components. The contents of photo-selective neoxanthin and lutein were high as well. Yet, β-carotene content was extraordinarily low. In addition, twigs retained high pre-dawn ratios of the deepoxidized antheraxanthin and zeaxanthin, although environmental conditions were not pre-disposing for such a state. The unexpected Car composition allows the conclusion that other micro-environmental conditions within twigs (hypoxia, increased red to blue photon ratios, and extremely high CO₂ concentrations) are more important than shade in shaping the Car profiles.     

Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis

The photosynthetic pigments of twigs in five tree and shrub species possessing chlorenchyma under a well developed, stomata-less, and highly photon absorptive periderm were analysed and compared to those of the corresponding canopy leaves. We asked whether the unavoidable shade acclimation of corticular chlorenchyma results in photosynthetic pigment complements typically found in shade leaves. As expected, chlorophyll (Chl) a/b ratios in twigs were consistently low. However, carotenoid (Car) analysis did not confirm the initial hypothesis, since twigs generally contained increased Chl-based pool sizes of the xanthophyll cycle components. The contents of photo-selective neoxanthin and lutein were high as well. Yet, -carotene content was extraordinarily low. In addition, twigs retained high pre-dawn ratios of the deepoxidized antheraxanthin and zeaxanthin, although environmental conditions were not pre-disposing for such a state. The unexpected Car composition allows the conclusion that other micro-environmental conditions within twigs (hypoxia, increased red to blue photon ratios, and extremely high CO₂ concentrations) are more important than shade in shaping the Car profiles.This revised version was published online in March 2005 with corrections to the page numbers.     

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     

Characterization of the Xanthophyll Cycle and Other Photosynthetic Pigment Changes Induced by Iron Deficiency in Sugar Beet (Beta vulgaris L.)

In this work we characterize the changes induced by iron deficiency in the pigment composition of sugar beet (Beta vulgaris L.) leaves. When sugar beet plants were grown hydroponically under limited iron supply, neoxanthin and β-carotene decreased concomitantly with chlorophyll a, whereas lutein and the carotenoids within the xanthophyll cycle were less affected. Iron deficiency caused major increases in the lutein/chlorophyll a and xanthophyll cycle pigments/chlorophyll a molar ratios. Xanthophyll cycle carotenoids in Fe-deficient plants underwent epoxidations and de-epoxidations in response to ambient light conditions. In dark adapted Fe-deficient plants most of the xanthophyll cycle pigment pool was in the epoxidated form violaxanthin. We show, both by HPLC and by in vivo 505 nanometers absorbance changes, that in Fe deficient plants and in response to light, the de-epoxidated forms antheraxanthin and zeaxanthin were rapidly formed at the expense of violaxanthin. Several hours after returning to dark, the xanthophyll cycle was shifted again toward violaxanthin. The ratio of variable to maximum chlorophyll fluorescence from intact leaves was decreased by iron deficiency. However, in iron deficient leaves this ratio was little affected by light conditions which displace the xanthophyll cycle toward epoxidation or de-epoxidation. This suggests that the functioning of the xanthophyll cycle is not necessarily linked to protection against excess light input.     

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.     

Carotenoid composition in sun and shade leaves of plants with different life forms

The carotenoid composition of sun leaves of nine species of annual crop plants (some with several varieties) was compared with sun and shade leaves of several other groups of plants, among those sun and shade leaves of several species of perennial shrubs and vines and deep-shade leaves of seven rainforest species. All sun leaves contained considerably greater amounts of the components of the xanthophyll cycle violaxanthin, antheraxanthin and zeaxanthin as well as of β-carotene than the shade leaves, as had previously been reported for a variety of other species by Thayer & Björkman (Photosynthesis Research, 1990, 23, 331–343). Therefore, high light specifically stimulated β,β-carotenoid synthesis. The sun leaves of these crop species did not contain α-carotene which was, however, present in large amounts in all shade leaves and in smaller amounts in sun leaves of three of the four species of perennial shrubs and vines. There was no difference in neoxanthin content on a chlorophyll basis between sun and shade leaves, and there was no consistent general difference in the lutein content between all sun and all shade leaves. The zeaxanthin (and antheraxanthin) content at peak irradiance and the xanthophyll cycle pool size were compared for sun leaves from the different groups of plants with different life forms and different metabolic activities. When growing in full sunlight the annual crop species and a perennial mesophyte had high rates of photosynthesis whereas the perennial shrubs and vines had relatively low photosynthesis rates. More zeaxanthin (and antheraxanthin) were accumulated at noon in full sunlight in those species with the lower photosynthesis rates. However, it was not such that those species also possessed the larger pools of violaxanthin + antheraxanthin + zeaxanthin. Instead, the xanthophyll cycle pools of sun leaves of the annual crop species and the perennial mesophyte were not smaller, and were even possibly larger, than those of sun leaves of the perennial shrubs and vines with low photosynthesis rates. This was so in spite of the fact that the crop species experienced much lesser degrees of excessive light at full sun than the shrubs and vines. Thus, many of the crop species converted only about 30–50% of their xanthophyll cycle pool to zeaxanthin at noon, whereas the shrubs and vines typically converted more than 80% of their pool into zeaxanthin. The crop species also had larger pools of β-carotene than the shrubs and vines but smaller pools of lutein than the majority of the latter species.     

Xanthophyll cycle components and capacity for non-radiative energy dissipation in sun and shade leaves ofLigustrum ovalifolium exposed to conditions limiting photosynthesis

The relationships between photosynthetic efficiency, non-radiative energy dissipation and carotenoid composition were studied in leaves ofLigustrum ovalifolium developed either under full sunlight or in the shade. Sun leaves contained a much greater pool of xanthophyll cycle components than shade leaves. The rate of non-radiative energy dissipation, measured as non-photochemical fluorescence quenching (NPQ), was strictly related to the deepoxidation state (DPS) of xanthophyll cycle components in both sun and shade leaves, indicating that zeaxanthin (Z) and antheraxanthin (A) are involved in the development of NPQ. Under extreme conditions of excessive energy, sun leaves showed higher maximum DPS than shade leaves. Therefore, sun leaves contained not only a greater pool of xanthophyll cycle components but also a higher proportion of violaxanthin (V) actually photoconvertible to A and Z, compared to shade leaves. Both these effects contributed to the higher NPQ in sun versus shade leaves. The amount of photoconvertible V was strongly related to chla/b ratio and inversely to leaf neoxanthin content. This evidence indicates that the amount of photoconvertible V may be dependent on the degree of thylakoid membrane appression and on the organization of chlorophyll-protein complexes, and possible explanations are discussed. Exposure to chilling temperatures caused a strong decline in the photon yield of photosynthesis and in the intrinsic efficiency of PS II photochemistry in sun leaves, but little effects in shade leaves. These effects were accompanied by increases in the pool of xanthophyll cycle components and in DPS, more pronounced in sun than in shade leaves. This corroborates the view that Z and A may play a photoprotective role under unfavorable conditions. In addition to the xanthophyll-related non-radiative energy dissipation, a slow relaxing component of NPQ, independent from A and Z concentrations, has been found in leaves exposed to low temperature and high light. This quenching component may be attributed either to other regulatory mechanism of PS II efficiency or to photoinactivation.Research supported by National Research Council of Italy, Special Project RAISA, Sub-Project 2, Paper N. 1587.     

Xanthophyll cycle pool size and composition in relation to the nitrogen content of apple leaves

The objective of this study was to determine xanthophyll cycle pool size and composition in response to N status and their relationships to non-photochemical quenching in apple leaves. Bench-grafted Fuji/M.26 trees were fertilized with different N concentrations (0-20 mM) in a modified Hoagland's solution for 6 weeks to create a wide range of leaf N status (1-4.4 g m(-2)). Chlorophyll content, xanthophyll cycle pool size, lutein, total carotene, and neoxanthin on a leaf area basis all increased linearly with increasing leaf N. However, only the ratios of the xanthophyll cycle pool and of lutein to chlorophyll were higher in low N leaves than in high N leaves. Under high light at midday, both zeaxanthin (Z), expressed on a chlorophyll basis, and the percentage of the xanthophyll cycle pool present as Z, increased as leaf N decreased. Thermal dissipation of excitation energy, measured as non-photochemical quenching of chlorophyll fluorescence, was positively related to, whereas efficiency of excitation transfer and photosystem II quantum efficiency were negatively related to, Z, expressed on a chlorophyll basis or on a xanthophyll cycle pool basis. It is concluded that both xanthophyll cycle pool size (on a chlorophyll basis) and conversion of violaxanthin to zeaxanthin are enhanced in response to N limitation to dissipate excessive absorbed light under high irradiance.     

Spectral Radiation Dependent Photoprotective Mechanism in the Diatom Pseudo-nitzschia multistriata

Phytoplankton, such as diatoms, experience great variations of photon flux density (PFD) and light spectrum along the marine water column. Diatoms have developed some rapidly-regulated photoprotective mechanisms, such as the xanthophyll cycle activation (XC) and the non-photochemical chlorophyll fluorescence quenching (NPQ), to protect themselves from photooxidative damages caused by excess PFD. In this study, we investigate the role of blue fluence rate in combination with red radiation in shaping photoacclimative and protective responses in the coastal diatom Pseudo-nitzschia multistriata. This diatom was acclimated to four spectral light conditions (blue, red, blue-red, blue-red-green), each of them provided with low and high PFD. Our results reveal that the increase in the XC pool size and the amplitude of NPQ is determined by the blue fluence rate experienced by cells, while cells require sensing red radiation to allow the development of these processes. Variations in the light spectrum and in the blue versus red radiation modulate either the photoprotective capacity, such as the activation of the diadinoxanthin-diatoxanthin xanthophyll cycle, the diadinoxanthin de-epoxidation rate and the capacity of non-photochemical quenching, or the pigment composition of this diatom. We propose that spectral composition of light has a key role on the ability of diatoms to finely balance light harvesting and photoprotective capacity.