二硫苏糖醇处理导致大豆叶片两光系统间激发能分配失衡

通过叶绿素荧光技术研究了二硫苏糖醇(1,4-dithiothreitol, DTT)对大豆叶片光系统I(PSI)和光系统Ⅱ(PSⅡ)间激发能分配的影响.结果显示:DTT处理没有影响叶片最大光化学效率(Fv/Fm),但光下叶绿素荧光降低比率(Rfd)下降;强光下,DTT处理叶片PSⅡ开放反应中心激发能捕获效率(Fv′/Fm′)比对照高30%～40%;分配给PSⅠ的激发能比对照叶片低约30%,分配给PSⅡ的激发能比对照叶片高20%左右,激发能分配严重偏离平衡状态;DTT处理叶片PSⅡ的激发能压力(1-qP)较对照高,但非光化学猝灭(qN)明显比对照低;进一步的实验揭示DTT的引入抑制了玉米黄质(Z)的生成和状态转换(qT).据此,推测DTT可能通过抑制天线色素的调节能力导致两光系统间激发能分配失衡.     

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     

缺锰降低大豆叶片叶绿素荧光的高能态猝灭

缺锰大豆叶片光合速率比对照低 5 0 %左右。缺锰叶片的Fv/Fm较对照低约 30 %～ 40 % ,而Fo比对照高 2～ 3倍左右。强光下 ,缺锰叶片的qP为对照的5 0 %。缺锰降低了叶片的高能态猝灭 (qE) ,仅占其NPQ的 6 0 %左右 ;而对照 qE组分占NPQ的 90 %左右。对照和缺锰叶片叶黄素库的脱环氧化程度分别为36 %和 2 1%左右。认为缺锰叶片 qE的降低可能是由于叶黄素脱环氧化程度减少所致     

Photoprotective Energy Dissipation in Higher Plant Leaves Investigated by Chlorophyll Fluorescence Decay Measurements with Additional Radiation Pulses

Chlorophyll (Chl) fluorescence decay measurements were performed on higher plant leaves to investigate the photoprotective mechanisms under in vivo conditions. Measurements on leaves with different amounts of zeaxanthin pointed out that zeaxanthin is necessary for most of the observed nonphotochemical energy quenching, that has to be activated by a transthylakoid pH. An additional sustained energy quenching component was clearly resolved in leaves with high amounts of zeaxanthin. The changes of the Chl fluorescence decay parameters did not correlate with a photoprotective energy dissipation in the reaction centre of photosystem 2 (P680), nor with a pH-mediated, zeaxanthin-independent aggregation of the antenna complexes; no indications for a state 1/state 2 transition of the main light-harvesting complex LHC2 were found.     

Photosynthesis of Hedera Canariensis var. Azorica Variegated Leaves as Affected by Ozone

A differential response to long-term ozone exposures (50 and 100 mm3 m-3) was observed in the green and white areas of variegated leaves of Hedera canariensis var. azorica L. In green tissue the photosynthetic activity was depressed via a stomatal mechanism, and in white regions no effect was observed. Chlorophyll fluorescence parameters remained unchanged in green portions, whereas in the white ones Fm and Fv/Fm significantly diminished following ozone fumigation.     

Alteration of Components of Chlorophyll-Protein Complexes and Distribution of Excitation Energy Between the Two Photosystems in Two New Rice Chlorophyll b-less mutants

Two new yellow rice chlorophyll (Chl) b-less (lack) mutants VG28-1 and VG30-5 differ from the other known Chl b-less mutants with larger amounts of soluble protein and ribulose-1,5-bisphosphate carboxylase/oxygenase small sub-unit and smaller amounts of Chl a. We investigated the altered features of Chl-protein complexes and excitation energy distribution in these two mutants, as compared with wild type (WT) rice cv. Zhonghua 11 by using native mild green gel electrophoresis and SDS-PAGE, and 77 K Chl fluorescence in the presence of Mg²⁺. WT rice revealed five pigment-protein bands and fourteen polypeptides in thylakoid membranes. Two Chl b-less mutants showed only CPI and CPa pigment bands, and contained no 25 and 26 kDa polypeptides, reduced amounts of the 21 kDa polypeptide, but increased quantities of 32, 33, 56, 66, and 19 kDa polypeptides. The enhanced absorption of CPI and CPa and the higher Chl fluorescence emission ratio of F685/F720 were also observed in these mutants. This suggested that the reduction or loss of the antenna LHC1 and LHC2 was compensated by an increment in core component and the capacity to harvest photon energy of photosystem (PS) 1 and PS2, as well as in the fraction of excitation energy distributed to PS2 in the two mutants. 77 K Chl fluorescence spectra of thylakoid membranes showed that the PS1 fluorescence emission was shifted from 730 nm in WT rice to 720 nm in the mutants. The regulation of Mg²⁺ to excitation energy distribution between the two photosystems was complicated. 10 mM Mg²⁺ did not affect noticeably the F685/F730 emission ratio of WT thylakoid membranes, but increased the ratio of F685/F720 in the two mutants due to a reduced emission at 685 nm as compared to that at 720 nm.     

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快、中和慢３ 个组分（ｑＮｆ,ｑＮｍ 与ｑＮｓ） 和５０５ ｎｍ 光吸收的日变化。主要结果如下：（１） 中午,珊瑚树叶片的ｑＮｓ 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ｑＮｓ 和ｑＮｆ 相比,除早晨和傍晚以外全天的水平都是最低的。（２） 珊瑚树叶片的初始荧光水平（Ｆｏ） 中午最低,而大豆叶片的Ｆｏ 中午最高。（３） 饱和光照射引起的珊瑚树叶片５０５ ｎｍ 光吸收的增加比大豆叶片大得多。（４） 珊瑚树叶片５０５ ｎｍ 光吸收的日变化方式与ｑＮｓ 的相类似。（５） 叶黄素循环的抑制剂ＤＴＴ对珊瑚树叶片ｑＮｓ 的抑制（５７ ％ ） 比对大豆叶片ｑＮｓ 的抑制（２３ ％ ） 严重。     

Dissipation of Excess Energy in Mehler-Peroxidase Reaction in Rumex Leaves During Salt Shock

By measurement of gas exchange and chlorophyll fluorescence, the effects of salt shock on photosynthesis and the mechanisms to protect photosynthetic machinery against photodamage during salt shock were investigated in leaves of Rumex seedlings. Salt shock induced significant decrease in photosynthesis both in 21 and 2 % O₂. In 21 % O₂, quantum yield of photosystem 2 (PS2) electron transport (_PS2) decreased slightly and q_P remained constant, suggesting that the excitation pressure on PS2 did not increase during salt shock. In 2 % O₂, however, both _PS2 and q_P decreased significantly, suggesting that the excitation pressure on PS2 increased during salt shock. NPQ increased slightly in 21 % O₂ whereas it increased significantly in 2 % O₂. The data demonstrated that during salt shock a considerable electron flow was allocated to oxygen reduction in the Mehler-peroxidase reaction (MPR). Under high irradiance and in the presence of saturating CO₂, the susceptibility of PS2 to photoinhibition in salt-shocked leaves was increased when the electron flow to oxygen in MPR was inhibited in 2 % O₂. Hence, MPR is important in photoprotection of Rumex seedlings during salt shock.     

Photosynthetic Pigments and Gas Exchange of in vitro Grown Tobacco Plants as Affected by CO2 Supply

Contents and functioning of photosynthetic pigments and gas exchange of Nicotiana tabacum L. leaves were studied in platlets cultivated in vitro under different CO₂ supply. The plantlets were grown for six weeks either in glass vessels tightly closed with aluminium foil (G-plants) or in polycarbonate Magenta GA-7 vessels covered with closures with microporous vents (M-plants). M-plants (better supplied with CO₂) had higher contents of chlorophyll (Chl) a. Chl b. and β-carotene, higher photochemical activities of photosystem 2 and whole electron transport chain, and lower contents of xanthophyll cycle pigments. Differences in Chl a fluorescence kinetic parameters between G-plants and M-plants were not statistically significant. M-plants had higher net photosynthetic rate, and lower transpiration rate and stomatal conductance than G-plants. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Characteristics of Photosynthetic Apparatus in Mn-Starved Maize Leaves

The effects of Mn-deficiency on CO₂ assimilation and excitation energy distribution were studied using Mn-starved maize leaves. Mn-deficiency caused about 70 % loss in the photon-saturated net photosynthetic rate (P _N) compared to control leaves. The loss of P _N was associated with a strong decrease in the activity of oxygen evolution complex (OEC) and the linear electron transport driven by photosystem 2 (PS2) in Mn-deficienct leaves. The photochemical quenching of PS2 (q_P) and the maximum efficiency of PS2 photochemistry (F_v/F_m) decreased significantly in Mn-starved leaves under high irradiance, implicating that serious photoinhibition took place. However, the high-energy fluorescence quenching (q_E) decreased, which was associated with xanthophyll cycle. The results showed that the pool of de-epoxidation components of the xanthophyll cycle was lowered markedly owing to Mn deficiency. Linear electron transport driven by PS2 de-creased significantly and was approximately 70 % lower in Mn-deficient leaves than that in control, indicating less trans-thylakoid pH gradient was built in Mn deficient leaves. We suggest that the decrease of non-radiative dissipation depending on xanthophyll cycle in Mn-starved leaves is a result of the deficiency of trans-thylakoid pH gradient.     

Influence of Manganese Toxicity on Photosynthesis in Ricebean (Vigna Umbellata) Seedlings

Influence of manganese (Mn) toxicity on photosynthesis in ricebean (Vigna umbellata) was studied by the measurement of gas exchange characteristics and chlorophyll fluorescence parameters. The net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) were reduced with increasing Mn concentration in nutrient solution. The reduction in g _s and E was more pronounced at 6 d of Mn treatment. However, P _N declined at 2 d of Mn treatment implying that the reduction in photosynthesis was not due to the direct effect of Mn on stomatal regulation. Mn did not affect the maximum efficiency of photosystem 2 (PS2) photochemistry (F_v/F_m). A reduction in photochemical quenching (q_P) and excitation capture efficiency of open PS2 (F_v′/F_m′) with a concomitant increase in q_N was observed. This implies that reduced demand for ATP and NADPH due to the reduction in photosynthesis causes a down-regulation of PS2 photochemistry and thus a high pH gradient (increase in q_N) and limited electron transport (decreased q_P). This revised version was published online in August 2006 with corrections to the Cover Date.     

田间大豆叶片成长过程中的光合特性及光破坏防御机制

田间大豆叶片在成长进程中光饱和光合速率持续提高,但气孔导度的增加明显滞后.尽管叶片在成长初期就具有较高的最大光化学效率,但是仍略低于发育成熟的叶片.随着叶片的成长,光下叶片光系统Ⅱ实际效率增加;非光化学猝灭下降.幼叶叶黄素总量与叶绿素之比较高,随着叶面积的增加该比值下降,在光下,幼叶的脱环氧化程度较高.因此认为大豆叶片成长初期就能够有效地进行光化学调节;在叶片生长过程中依赖叶黄素循环的热耗散机制迅速建立起来有效抵御强光的破坏.     

Photosynthesis and photoprotection in Nicotiana tabacum L. in vitro-grown plantlets

Nicotiana tabacum L. plantlets were cultured in vitro photoautotrophically (0% sucrose) and photomixotrophically (3% or 5% sucrose) at two irradiances (80 or 380 mumol m-2 s-1) with the aim of investigating the effect of these culture conditions on photosynthetic parameters and on protective systems against excess excitation energy. In plantlets grown photoautotrophically under higher irradiance photoinhibition was demonstrated. These plantlets had a decreased chlorophyll (Chl) a + b content and Chl a/b ratio, an increased content of xanthrophyll cycle pigments and a higher deepoxidation state, a decreased maximum photochemical efficiency of photosystem II (PS II) and actual photochemical efficiency of PS II, and an increased non-photochemical quenching. In the photoautotrophically grown plantlets and those photomixotrophically grown with 3% sucrose, the increase of growth irradiance from 80 to 380 mumol m-2 s-1 stimulated the activities of ascorbate-glutathione cycle enzymes with the exception of ascorbate peroxidase. Ascorbate peroxidase activity was not affected by the increase in growth irradiance but a significant decrease with increasing sucrose concentration was evident. The higher concentration of sucrose in the medium (5%) in combination with the higher irradiance inhibited photosynthesis (decrease in Chl a + b content and net photosynthetic rate) but no significant changes in activities of ascorbate-glutathione cycle enzymes were found. These results suggest that exogenous sucrose added to the medium improved high irradiance and oxidative stress resistance of the plantlets but the effect of sucrose is concentration dependent.     

Regulation Mechanism of Excitation Energy Transfer in Phycobilisome-Thylakoid Membrane Complexes

Regulation mechanism of excitation energy transfer between phycobilisomes (PBS) and the photosynthetic reaction centres was studied by the state transition techniques in PBS-thylakoid membrane complexes. DCMU, betaine, and N-ethylmaleimide were applied to search for the details of energy transfer properties based on the steady fluorescence measurement and individual deconvolution spectra at state 2 or state 1. The closure of photosystem (PS) 2 did not influence on fluorescence yields of PS1, i.e., energy could not spill to PS1 from PS2. When the energy transfer pathway from PBS to PS1 was disturbed, the relative fluorescence yield of PS2 was almost the same as that of PS2 in complexes without treatment. If PBSs were fixed by betaine, the state transition process was restrained. Hence PBS may detach from PS2 and become associated to PS1 at state 2. Our results contradict the proposed "spill-over" or "PBS detachment" models and support the mobile "PBS model".     

Different Relationship Between Electron Transport and CO2 Assimilation in two Zea mays Cultivars as Influenced by Increasing Irradiance

Gas exchange and fluorescence parameters were measured simultaneously in two Zea mays L. cultivars (Liri and 121C D8) to assess the relationship between the quantum yield of electron transport (_PS2) and the quantum yield of CO₂ assimilation (_CO2) in response to photosynthetic photon flux density (PPFD). The cv. Liri was grown under controlled environmental conditions in a climate chamber (CC) while cv. 121C D8 was grown in CC as well as outdoors (OT). By exposing the two maize cultivars grown in CC to an increasing PPFD, higher photosynthetic and photochemical rates were evidenced in cv. Liri than in cv. 121C D8. In Liri plants the _PS2/_CO2 ratio increased progressively up to 27 with increasing PPFD. This suggests that the reductive power was more utilised in non-assimilatory processes than in CO₂ assimilation at high PPFD. On the contrary, by exposing 121C D8 plants to increasing PPFD, _PS2/_CO2 was fairly constant (around 11–13), indicating that the electron transport rate was tightly down regulated by CO₂ assimilation. Although no significant differences were found between _PS2/_CO2 of the 121C D8 maize grown under CC and OT by exposing them to high PPFD, the photosynthetic rate and photochemical rates were higher in OT maize plants.     

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.     

Photosynthesis in different types of transgenic tobacco plants with elevated cytokinin content

Photosynthetic parameters were compared in three types of transgenic tobacco plants: ipt-transgenic plants with slightly elevated endogenous cytokinin (CK) content, Pssu-ipt-transgenic plants with markedly increased CK content, and zmp-transgenic plants with slightly elevated CK content accompanied by elevated auxin content. Slightly increased CK content promoted net photosynthetic rate (P_N) in both ipt- and zmp-transgenic plants, and chlorophyll (Chl) and carotenoid contents in zmp-transgenic plants. Morphology, growth characteristics, stomatal conductance, and parameters of Chl a fluorescence kinetics were similar in control and transgenic plants with slightly higher CK content. No significant effect of increased level of endogenous auxin (indole-3-acetic acid) on development of zmp-transgenic plants and measured parameters was found. Pssu-ipt-transgenic plants with highly increased CK content revealed suppressed root development, wilting of plants, and depression of P_N and stomatal conductance; however, Chl content was slightly increased and parameters of Chl a fluorescence kinetics did not indicate damage to photosynthetic apparatus. This revised version was published online in July 2006 with corrections to the Cover Date.     

青海薄荷的光合及热能耗散特性研究

对青海省民和、西宁和湟源3个地区的药用植物薄荷(Mentha haplocalyx Briq.)的净光合速率和叶绿素荧光参数的变化进行分析比较。结果显示:(1)最大净光合速率(Pn)湟源>西宁>民和;(2)PSⅡ反应中心最大光化学效率(Fv/Fm)以及PSⅡ反应中心活性参数(1/Fo-1/Fm)和实际光化学量子效率(ΦPSⅡ)的变化趋势相同,均以西宁最高,而相应的光合功能的相对限制(LPDF)西宁最低;(3)西宁薄荷的光化学猝灭系数(qP)最高(0.875),而非光化学猝灭(NPQ)则最低(1.114)。研究表明,青海薄荷有光抑制现象发生,但并没有造成PSⅡ反应中心的不可逆破坏;西宁薄荷光化学能力较高而湟源薄荷对于过剩光能的耗散能力较高。     

Protective Action of Abscisic Acid Against the Inhibition of Photosynthesis of Barley Leaves by Bisulphite

The inhibition of photosynthetic activity by bisulphite was studied in intact leaves of abscisic acid (ABA)-treated and non-treated (control) barley plants. ABA inhibited the photosynthetic process as evidenced by lower values of chlorophyll fluorescence kinetic parameters F_v/F_m (photosystem 2 activity) and R_fd (vitality index, related to the whole photosynthetic activity) compared with ABA-non-treated plants. After bisulphite treatment, the extent of inhibition was smaller in ABA-treated plants than in the control ones indicating a protective effect of ABA. The protective action sites of ABA were the Q_A reduction and the Calvin cycle. This revised version was published online in September 2006 with corrections to the Cover Date.