Enhanced Thermal Energy Dissipation Depending on Xanthophyll Cycle and D1 Protein Turnover in Iron-Deficient Maize Leaves Under High Irradiance

Pigment contents of chloroplasts and net photosynthetic rate were dramatically reduced in maize leaves suffering from iron deficiency. However, the reduction in photosynthesis was probably not caused by decreased contents of chlorophylls and carotenoids and by photon absorption; the primary limiting factor for photosynthesis may rather be the decrease of electron transport activity in photosystem 1. Iron-deficient leaves suffered serious acceptor-side photoinhibition, and more than 60 % of absorbed photons were dissipated, while less than 40 % was used in photochemical reaction. Thermal energy dissipation depending on xanthophyll cycle and D1 protein turnover was enhanced when acceptor-side photoinhibition occurred in iron-deficient maize leaves.     

Xanthophyll cycle activity in detached barley leaves senescing under dark and light

Senescence-induced changes in the xanthophyll cycle activity and chlorophyll (Chl) fluorescence parameters were compared in detached barley (Hordeum vulgare L.) leaf segments kept for 6 d in darkness or under continuous white light (90 mol m^–2 s^–1). Before detachment of the leaf segments, the plants were grown at periodic regime [12 h light (90 mol m^–2 s^–1)/12 h dark]. The de-epoxidation state of the xanthophyll cycle pigments (DEPS) in the leaf samples was determined immediately (the actual DEPS), after 1 h of dark-adaptation (the residual DEPS), and during 14 min of a high-irradiance (HI) exposure (500 mol m^–2 s^–1) (HI-induced DEPS). In the light-senescing segments, senescence was delayed pronouncedly compared to dark-senescing ones as the Chl content, the photosystem 2 photochemistry, and electron transport processes were highly maintained. Further, the actual DEPS increased, probably due to the increased mean photon dose. The HI-induced increase in the DEPS was stimulated in the light-senescing segments, whereas it was slowed down in the dark-senescing ones. However, after the 14 min HI-exposure of the dark-senescing segments the HI-induced DEPS was not markedly lower than in the mature leaves, which indicated the maintenance of the xanthophyll cycle operation.     

Photoinhibition Characteristics of a Low Chlorophyll b Mutant of High Yield Rice

The low chlorophyll b mutant of high yield rice had a lower light-harvesting complex 2 content than the wild type. The stability of oxygen evolution side of photosystem 2 was only slightly lower. A lower photon absorption rate and a stronger xanthophyll cycle capacity of this mutant led to a higher endurance to strong irradiance and a lower photoinhibition as compared with the wild type rice.     

The Effect of Low Growth Temperature on Hill Reaction and Photosystem 1 Activities and Pigment Contents in Maize Inbred Lines and Their F1 Hybrids

Young plants of maize inbred lines CE777, CE704, and CE810 and their F₁ hybrids displaying a positive heterotic effect in various photosynthetic characteristics were exposed to low temperature during their early growth developmental stage. The photochemical activity of isolated mesophyll chloroplasts and the contents of photosynthetic pigments in leaves of stressed and non-stressed plants were compared with the aim to find out the possible changes in the relationship between parents and hybrids, and to determine the genetic basis of heterosis in F₁ generation. Strong decrease in the content of chlorophylls was observed for all genotypes examined when plants were subjected to low growth temperature. Similar change was recorded for Hill reaction activity (HRA) of inbred lines but not of their F₁ hybrids, and no significant response at all was found for photosystem 1 (PS1) activity or the total carotenoids content. The intraspecific variation due to differences between genotypes was found for most of photosynthetic characteristics examined. This variation was caused by the additive and dominance genetic effects. Positive dominance was the main cause of positive heterosis in HRA and in the contents of photosynthetic pigments and was much more pronounced in the stressed plants compared to the non-stressed ones. The maternal additive effects participated in the inheritance of contents of photosynthetic pigments in plants exposed to low temperature, too. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of Cold-Hardening on Chilling-Induced Photoinhibition of Photosynthesis and on Xanthophyll Cycle Pigments in Sweet Pepper

Two cultivars of Capsicum annuum L. were acclimated for 5 d at sub-optimal temperature (14 °C) and irradiance of 250 µmol m^–2 s^–1. This cold-hardening resulted in some reduction in the extent of photoinhibition during an 8 h exposure to high irradiance at 4 °C. Obvious differences were observed between non-hardened leaves (NHL) and cold-hardened leaves (CHL) in the recovery under low irradiance at room temperature. The CHL of both cultivars recovered faster than NHL, especially during the initial fast phase of recovery. Compared with NHL, the total content of carotenoids (Cars), based on chlorophyll, Chl (a+b), and the proportions of xanthophyll cycle pigments referred to total Cars increased in CHL, mainly due to an increase of violaxanthin (V) + antheraxanthin (A) + zeaxanthin (Z) content per mol Chl (a+b). Faster development and a higher non-photochemical quenching (NPQ) of Chl fluorescence, related to a stronger deepoxidation of the larger xanthophyll cycle pool in NHL, could act as a major defence mechanism to reduce the formation of reactive oxygen species during severe chilling. This is suggested by higher content of Z or Z+A in photoinhibition as well as by its rapid decline during the initial fast phase of recovery. In contrast to the chilling-sensitive cv. 0004, the chilling-tolerant cv. 1141 did more easily acclimate its photosynthetic apparatus to low temperatures.     

Photoinhibition and Active Oxygen Species Production in Detached Apple Leaves During Dehydration

In the course of dehydration, the gas exchange and chlorophyll (Chl) fluorescence were measured under irradiance of 800 mol m^–2 s^–1 in detached apple leaves, and the production of active oxygen species (AOS), hydrogen peroxide (H₂O₂), superoxide (O₂ ^–), hydroxyl radical (–OH), and singlet oxygen (¹O₂), were determined. Leaf net photosynthetic rate (P _N) was limited by stomatal and non-stomatal factors at slight (2–3 h dehydration) and moderate (4–5 h dehydration) water deficiency, respectively. Photoinhibition occurred after 3-h dehydration, which was defined by the decrease of photosystem 2 (PS2) non-cyclic electron transport (P-rate). After 2-h dehydration, an obvious rise in H₂O₂ production was found as a result of photorespiration rise. If photorespiration was inhibited by sodium bisulfite (NaHSO₃), the rate of post-irradiation transient increase in Chl fluorescence (Rfp) was enhanced in parallel with a slight decline in P-rate and with an increase in Mehler reaction. At 3-h dehydration, leaf P-rate decrease could be blocked by glycine (Gly) or methyl viologen (MV) pre-treatment, and MV was more effective than Gly at moderate drought time. AOS (H₂O₂ and O₂ ^–), prior to photoinhibition produced from photorespiration and Mehler reaction in detached apple leaves at slight water deficiency, were important in dissipating photon energy which was excess to the demand of CO₂ assimilation. So photoinhibition could be effectively prevented by the way of AOS production.     

Saturating Irradiance-Induced Photoinhibition without Monomerisation of Photosystem 2 Dimer in Soybean Leaves

The oligomeric state of photosystem 2 (PS2) complex in soybean leaves treated with saturating irradiance was studied by non-denaturing polyacrylamide gel electrophoresis (PAGE) and gel filtration chromatography. PS2 dimers resolved by non-denaturing PAGE accounted for about 75 % of total PS2 complex and there was no significant difference in the ratio of PS2 dimer to monomer between samples from saturating irradiance-treated and fully dark-adapted leaves. Furthermore, BBY particles were resolved into four chlorophyll-enriched fractions by gel filtration chromatography. From their molecular masses and protein components, these fractions were deduced to be PS2 dimer, PS2 monomer, oligomeric light-harvesting complex 2 (LHC2), and monomeric LHC2. Also, no change in the proportion of PS2 dimer in total PS2 was observed in the granal region of thylakoid membranes from soybean leaves after saturating irradiation. Hence the dimer is the predominant natural form of PS2 in vivo and no monomerisation of PS2 dimer occurs during saturating irradiance-induced photoinhibition in soybean leaves.     

Synergistic Effect of AlCl3 and Kinetin on Chlorophyll and Protein Contents and Photochemical Activities in Detached Wheat Primary Leaves during Dark Incubation

Al³⁺ in combination with kinetin showed more protection against degradation of chlorophyll (Chl) and protein than Al³⁺ or kinetin alone during dark-induced senescence in wheat primary leaf segments. MV-dependent whole chain electron transport, photosystem (PS) 2 mediated oxygen evolution, and PS1 activities were also delayed to a greater extent. Absorbed excitation energy distribution was more in favour of PS1 in Al³⁺ plus kinetin-treated leaf thylakoids at 72 h.     

Relationship Between Photosystem 2 Electron Transport and Photosynthetic CO2 Assimilation Responses to Irradiance in Young Apple Tree Leaves

The responses to irradiance of photosynthetic CO₂ assimilation and photosystem 2 (PS2) electron transport were simultaneously studied by gas exchange and chlorophyll (Chl) fluorescence measurement in two-year-old apple tree leaves (Malus pumila Mill. cv. Tengmu No.1/Malus hupehensis Rehd). Net photosynthetic rate (P _N) was saturated at photosynthetic photon flux density (PPFD) 600-1 100 (mol m^-2 s^-1, while the PS2 non-cyclic electron transport (P-rate) showed a maximum at PPFD 800 mol m^-2 s^-1. With PPFD increasing, either leaf potential photosynthetic CO₂ assimilation activity (F_d/F_s) and PS2 maximal photochemical activity (F_v/F_m) decreased or the ratio of the inactive PS2 reaction centres (RC) [(F_i – F_o)/(F_m – F_o)] and the slow relaxing non-photochemical Chl fluorescence quenching (q_s) increased from PPFD 1 200 mol m^-2 s^-1, but cyclic electron transport around photosystem 1 (RFp), irradiance induced PS2 RC closure [(F_s – F_o)/F_m – F_o)], and the fast and medium relaxing non-photochemical Chl fluorescence quenching (q_f and q_m) increased remarkably from PPFD 900 (mol m^-2 s^-1. Hence leaf photosynthesis of young apple leaves saturated at PPFD 800 mol m^-2 s^-1 and photoinhibition occurred above PPFD 900 mol m^-2 s^-1. During the photoinhibition at different irradiances, young apple tree leaves could dissipate excess photons mainly by energy quenching and state transition mechanisms at PPFD 900-1 100 mol m^-2 s^-1, but photosynthetic apparatus damage was unavoidable from PPFD 1 200 mol m^-2 s^-1. We propose that Chl fluorescence parameter P-rate is superior to the gas exchange parameter P _N and the Chl fluorescence parameter F_v/F_m as a definition of saturation irradiance and photoinhibition of plant leaves.     

干旱条件下冬小麦不同叶龄叶绿素荧光及叶黄素循环组分的变化(英文)

干旱条件下冬小麦叶片光化学效率（Ｆｖ／Ｆｍ）的降低伴随着叶黄素循环组分玉米黄质含量的增加。干旱初期,Ｆｖ／Ｆｍ 的降低约在暗置过夜后完全恢复。当干旱诱导的玉米黄质含量的增加达最大值时,Ｆｖ／Ｆｍ 不可逆下降,Ｆｏ 上升,表明发生了光破坏。与老叶相比,干旱条件下功能叶具有较高的玉米黄质含量,对光破坏的抗性较强。推测干旱条件下老叶不可逆衰老与其依赖于叶黄素循环的耗散过剩光能能力的下降有关。     

低剂量UV-B辐射对乌拉尔甘草叶片光合机构的影响

为了探讨植物叶片对UV-B辐射增强的响应机制,采用叶绿素荧光测定技术,分别测定在人工模拟低剂量UV-B(2.4μW/cm~2)辐射条件下乌拉尔甘草叶片的叶绿素荧光诱导动力曲线、初始荧光(F_0)、最大荧光(F_m)、光合机构比活性参数(ABS/RC、TR_o/RC和ET_o/RC)和性能指数等变化规律。结果表明:(1)低剂量UV-B辐射未引起甘草叶片O-J-I-P叶绿素荧光诱导曲线中的相数发生改变,UV-B辐射对PSⅡ的影响主要发生在其受体侧,而非供体侧;(2)低剂量UV-B辐射引起了甘草叶片光合系统F_v/F_m以及F_m、F_0的明显变化,同时也影响了光合机构的开放程度和电子从Q_A向Q_B传递效率,从而影响了光转化效率;相应性能指数(PI_(abs)和PI_(total))的改变亦验证了此结果。研究认为,低剂量UV-B辐射抑制乌拉尔甘草叶片光合系统Ⅱ受体侧Q_A至PQ之间的电子传递效率,从而影响了Q_A之后的光化学反应及非光化学反应。     

Correlation between Photoinhibition Sensitivity and the Rates and Relative Extents of Xanthophyll Cycle De-epoxidation in Chlorina Mutants of Barley (Hordeum vulgare L.)

We compared photoinhibition sensitivity to high irradiance (HI) in wild-type barley (wt) and both its chlorina f ₁₀₄-nuclear gene mutant, that restricts chlorophyll (Chl) a and Chl b synthesis, and its f ₂-nuclear gene mutant, that inhibits all Chl b synthesis. Both F_v/F_m and _PS2 decreased more significantly in f ₂ than f ₁₀₄ and wt with duration of HI exposure. Chl degraded more rapidly in the f ₂ than in either f ₁₀₄ or wt. Most sensitivity to photoinhibition was exhibited for f ₂, whereas there was little difference in response to HI between the f ₁₀₄ and wt. The highest de-epoxidation (DES) value at every time point of exposure to HI was measured for f ₂, whereas the wt had the lowest value among the three strains. There were two lifetime components resolved for the conversion of violaxanthin (V) to zeaxanthin plus antheraxanthin (Z + A). The most rapid lifetime was around 6 min and the slower lifetime was >140 min, in both the mutants and wt. However, the wt and f ₁₀₄ both displayed larger amplitudes of both de-epoxidation lifetimes than f ₂. The difference between the final de-epoxidation state (DES = [Z + A]/[V + A + Z]) in the light compared to the dark expressed as DES for wt, f ₁₀₄, and f ₂ was 0.630, 0.623, and 0.420, respectively. The slow lifetime component and overall larger DES in the wt and f ₁₀₄ correlated with more photoprotection, as indicated by relatively higher F_v/F_m and _PS2, compared to the f ₂. Hence the photoprotection against photoinhibition has no relationship with the absolute DES value, but there is a strong relationship with de-epoxidation rate and relative extent or DES.     

Chlorophyll a Fluorescence Emission,Xanthophyll Cycle Activity,and Net Photosynthetic Rate Responses to Ozone in Some Foliose and Fruticose Lichen Species

The lichens Parmelia quercina, Parmelia sulcata, Evernia prunastri, Hypogymnia physodes, and Anaptychia ciliaris were exposed to ozone (O₃) in controlled environment cuvettes designed to maintain the lichens at optimal physiological activity during exposure. Measurements of gas exchange, modulated chlorophyll (Chl) fluorescence, and pigment analysis were conducted before and after exposure to 300 mm³ (O₃) m^–3, 4 h per d for 14 d. No changes in the efficiency of photosystem 2 (PS2) photochemistry, the reduction state of Q_A, or the electron flow through PS2, measured by Chl fluorescence, were detected in any of the five lichen species studied. Additionally, neither photosynthetic CO₂ assimilation nor xanthophyll cycle activity or photosynthetic pigment concentration were affected by high O₃ concentrations. Thus the studied lichen species have significant capacities to withstand oxidative stresses induced by high concentration of O₃.     

The Effect of In Vitro Culture Conditions on the Pattern of Photoinhibition during Acclimation of Gardenia Plantlets to Ex Vitro Conditions

We tested the effect of growing conditions during micropropagation on the fast kinetics of chlorophyll (Chl) fluorescence of Gardenia jasminoides Ellis plantlets during a 4-week acclimation to ex vitro. We studied whether photoautotrophic growing in vitro produced plantlets with less photoinhibition impairment during acclimation. Of the growing conditions stimulating photoautotrophy in vitro, only loose tube caps had a positive effect, whereas low sucrose or sucrose-free content in the medium and high PPFD showed a negative effect. Thus, plantlets cultured with 3 % (m/v) of sucrose were subsequently less photoinhibited throughout acclimation than those cultured with low sucrose (0.5 %) or sucrose-free media. Moreover, at the end of acclimation the former plantlets showed F_v/F_m and F_v/F₀ ratios typical of unstressed ex vitro plants as well as a higher Chl content and ratio of Chls to carotenoids. Plantlets cultured at a photosynthetic photon fluence density (PPFD) of 50 µmol m^–2 s^–1 also showed a better performance at the end of acclimation than those cultured at a higher (110 µmol m^–2 s^–1) PPFD. Thus except in the case of loose-tube closure, gardenia plantlets cultured in vitro under conventional sucrose concentration and PPFD are the least photoinhibited during acclimation. Nevertheless, significant interactions between the in vitro growing factors were observed at the end of acclimation.     

High Irradiance Effects on the Xanthophyll Cycle Pigments and the Activity of Violaxanthin De-Epoxidase in Soybean Callus

High irradiance (HI) effects on xanthophyll cycle pigments (XCP) and activity of violaxanthin de-epoxidase (VDE) in terms of de-epoxidation index (DEI) were studied in soybean calli. The calli from the hypocotyl segments of 5-d seedlings were induced on a solid (1.1 % agar) MS medium (pH 5.8) supplemented with 4.52 M 2,4-dichloro-phenoxyacetic acid, 2.32 M kinetin, and 3 % sucrose. After a 30 d cultivation, the green calli were irradiated for 24 h with white light (HI, 1 300 mol m^–2 s^–1) and VDE was isolated from the photosystem 2 (PS2) particles. In the control (0 h irradiation) callus, the reaction of PS2 particles with VDE in the presence or absence of Tween 20 resulted in the decrease of VIO content and the increase of ZEA content. In the 24 h HI-callus, the reaction of PS2 particles in the absence of VDE led to the decrease of VIO and ANT contents and increase of ZEA content. In the control, DEIs in the presence of VDE with or without 0.1 %Tween 20 (1.04 and 1.06, respectively) were significantly higher than the DEI (0.76) in the absence of VDE. In the HI-callus, DEIs in the presence of VDE with or without 0.1 %Tween 20 (0.98 and 0.96, respectively) were similar to that (1.03) in the absence of VDE.     

Comparative Characteristics of Growth and Photosynthesis of Sun and Shade Leaves from Normal and Pendulum Walnut (Juglans regia L.) Trees

Pendulum walnut leaves exhibited various adaptive responses related to the regulation of photon interception such as specific downward orientation, greater leaf area, and larger pigment pool. Changes in the regulation of PS2 such as higher thermal dissipation (NPQ) and lower quantum efficiency (Φ_PS2) that protect the photosynthetic apparatus against damages were also found. The growth and photosynthetic features of pendulum walnut leaf are interpreted as adaptations that allow the pendulum walnut tree to compensate the impaired ability by appropriate growth to ensure the energy needs for photosynthesis, respectively for biomass formation. This revised version was published online in August 2006 with corrections to the Cover Date.     

EGTA和酒石酸对蓖麻Cd胁迫与积累的调控作用

通过盆栽试验研究了解毒剂酒石酸与螯合剂EGTA的单施与配施对强化蓖麻修复Cd污染土壤的作用,探讨重金属污染土壤植物修复中螯合剂与解毒剂配合使用的可行性。结果显示:(1)除酒石酸单施处理外,其余处理均可显著提高土壤中醋酸提取态Cd含量,增强土壤Cd的活性,并以酒石酸与EGTA配施的效果更显好,其土壤醋酸提取态Cd含量为对照的1.41~2.49倍。(2)EGTA能有效促进Cd从蓖麻根部向地上部的转移,但高剂量EGTA处理对蓖麻根系有明显的毒害作用;EGTA与酒石酸配合施能缓解Cd对植株的毒害作用,增大蓖麻生物量和Cd积累量,其地上部Cd积累量比对照增加4.56~8.32倍。(3)蓖麻叶片Cd含量、地上部积累总量以及土壤净化率随土壤醋酸提取态Cd含量的升高而增大,并且呈良好的线性递增关系。研究表明,酒石酸与EGTA配施可通过调控土壤Cd的植物可利用性和降低Cd的生理毒性来提高蓖麻对Cd的富集能力和对Cd污染土壤的修复效果。     

Comparison of Aerial and Submerged Leaves in Two Amphibious Species,Myosotis scorpioides and Ranunculus trichophyllus

Both amphibious species, Myosotis scorpioides and Ranunculus trichophyllus, thrive in a stressful environment (alternated flooding and drying), which is variable regarding water and radiation regimes. Plants from the field and plants grown under controlled water table maintained at 40 cm were analysed for content of chlorophyll (Chl) and UV-B screening compounds, and the efficiencies of PS2 and electron transport systems. We detected no significant differences in contents of Chl and UV-B screening compounds between submerged and aerial leaves. The measurements of respiratory potential and photochemical efficiency revealed the presence of permanent stress in M. sporpioides in the natural environment. Differences in physiological responses of submerged and aerial leaves indicated that the terrestrial environment was more favourable for M. scorpioides than for R. trichophyllus. Characteristics of both species suggested that R. trichophyllus might be a phylogenetically older aquatic plant than M. scorpioides.     

Effect of Polyamines on Chlorophyll and Protein Contents, Photochemical Activity, and Energy Transfer in Detached Wheat Leaves during Dark Incubation

Spermine as compared to putrescine or spermidine retarded the loss of chlorophyll and protein contents to a greater extent in wheat primary leaves during dark incubation. Activities of whole chain electron transport, photosystem (PS) 1 and PS2, and absorbed excitation energy distribution in favour of PS 1 were protected by these amines in valency dependent manner during 72-h dark incubation.     

Photosynthetic Performance of Ginkgo biloba L. Grown Under High and Low Irradiance

Diurnal variation in net photosynthetic rate (P _N) of three-year-old plants of Ginkgo biloba was studied under open, O (receiving full sunlight), net-shade, NS (40 % of photosynthetically active radiation, PAR), or greenhouse, G (25 % PAR) conditions. In all three conditions, P _N was higher in morning along with stomatal conductance (g _s), and intercellular CO₂ concentration (C _i), while leaf temperature and vapour pressure deficit were low. The O-plants exhibited a typical decline in P _N during midday, which was not observed in NS-plants. This indicated a possible photoinhibition in O-plants as the ratio of variable to maximum fluorescence (F_v/F_m) and photosystem 2 (PS2) yield (_PS2) values were higher in the NS- and G-plants. On the contrary, stomatal density and index, chlorophyll a/b ratio, leaf thickness, and density of mesophyll cells were greater in O-plants. Further, higher P _N throughout the day along with higher relative growth rate under NS as compared to O and G suggested the better efficiency of Ginkgo plants under NS conditions. Therefore, this plant species could be grown at 40 % irradiance to meet the ever-increasing demand of leaf and also to increase its export potential.