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.     

温州蜜柑叶片光合作用光抑制的保护机理

晴天条件下,使用便携式调制荧光仪和分光光度计观察了温州蜜柑叶片光合作用光抑制发生过程中几个主要荧光参数（初始荧光F0、最大荧光Fm、PSⅡ的光化学效率Fv/Fm、非光化学猝灭qN及其快相qNf和慢相qNs）、电子传递速率（ETR）和玉米黄素相对含量的日变化,结果表明,随着光强的增强,ETR、qN及其qNr与qNs以及玉米黄素相对含量升高,Fv/Fm、Fm和F0下降。用DTT处理后,qNs较对照明显下降,F0较对照明显上升,可以认为,柑橘在光合作用日变化中存在依赖于叶黄素循环和类囊体膜质子梯度两种非辐射能量耗散方式,而且它们在防御光破坏方面起着重要作用。     

鸢尾(Iris L.)叶片取向与其光合特性及光抑制的关系

通过气体交换、叶绿素荧光、反射光谱等方法,研究了鸢尾叶片取向对植株光合特性及光抑制的影响.自然状态下,鸢尾的叶片不同取向影响植株对光能的截获;叶片净光合速率Pn与光合有效辐射PAR呈极显著相关;东西取向叶片的Pn要大于南北取向.南北取向的植株中叶片叶绿素(Chl a和Chl b),类胡萝卜素(Car)含量略高于东西取向.日进程中,各取向的叶片在一天中均没有发生明显的光抑制.相对于东西取向的植株,南北取向植株发生了明显的倾斜;在两种取向的植株中,叶片东侧和南侧的光化学反射指数(PRI)下调幅度较大;PRI的变化量(△PRI)大小依次为:东侧>南侧>西侧>北侧.鸢尾植株取向改变了叶片倾斜角度,两者共同导致光能截获减小;同时,叶片光能利用效率下调和叶黄素循环增强,这可能是不同取向植株均未发生严重光抑制的原因.     

棉花苞叶光呼吸和PSII热耗散对土壤水分的响应

在新疆气候生态条件下, 采用膜下滴灌植棉技术, 设置不同滴灌水分处理, 研究了不同滴灌量条件下棉花(Gossypium hirsutum)苞叶和叶片碳同化、光呼吸作用、光系统II (PSII)热耗散作用及其光破坏防御机制的差异, 以揭示滴灌节水条件下棉花苞叶缓解光抑制的机理及与棉花抗旱特性的关系。结果表明: 棉花开花后苞叶及叶片在高温强光下实际光化学效率(Φ_PSII)显著降低, 发生明显的光抑制现象, 但苞叶的光抑制程度较叶片轻; 与正常滴灌量处理相比, 节水滴灌条件下棉花水分亏缺, 叶片净光合速率(P_n)、Φ_PSII、光呼吸(P_r)、光化学猝灭系数(q_P)降低, 非光化学猝灭系数(NPQ)升高, 叶片光抑制程度加重, 而苞叶P_n、Φ_PSII、P_r、q_P、NPQ变化不大, 与正常滴灌量处理相比, 光抑制程度无显著差异。苞叶光呼吸速率与光合速率的比值(P_r/P_n)显著高于叶片; 滴灌节水条件下棉花适度水分亏缺对苞叶光呼吸及P_r/P_n无显著影响。高温强光下, 棉花节水滴灌对叶片PSII量子产量的转化与分配影响显著, 但对苞叶的影响不显著; 苞叶非调节性能量耗散的量子产量(Y(NPQ))高于叶片, 因此能有效地将PSII的过剩光能以热的形式耗散。综上所述, 与叶片相比, 苞叶对轻度水分亏缺不敏感, 是棉花适应干旱逆境较强的器官, 苞叶光呼吸和热耗散作用对光破坏防御具有重要意义。     

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。     

银杏叶片的光抑制和光保护机制：温度,CO2和O2的影响

田间条件下,夏季晴天中午银杏（ＧｉｎｋｇｏｂｉｌｏｂａＬ．）叶片经常遭受强光（超过１４００μｍｏｌ·ｍ－２·ｓ－１）和高温（３５℃左右）胁迫,气孔导度和ＣＯ２同化速率出现明显的“午休”现象。叶黄素循环关键组分玉米黄质在一天中随着光强而变化。在室内分析了强光和高温交互作用对银杏叶片光抑制的影响。在１５～３５℃温度范围内和１４００μｍｏｌ·ｍ－２·ｓ－１光强下,银杏叶片分别在空气中、低ＣＯ２浓度（８０μＬ·Ｌ－１）和２％Ｏ２条件下处理２ｈ。银杏叶肉ＣＯ２导度低（约３１ｍｍｏｌ·ｍ－２·ｓ－１）,导致羧化部位ＣＯ２浓度低,光呼吸较强。在空气中,电子传递与固定ＣＯ２之比较高（１６ｅ－／ＣＯ２,２５℃）。此比率在２％Ｏ２条件下仍随温度的升高而增大,仅用光呼吸的变化不能解释。在１５～３５℃温度范围内和不同ＣＯ２浓度下,降低Ｏ２虽然导致电子传递速率下降,但对光抑制程度无影响,表明强光条件下光呼吸对银杏叶片无保护作用。依赖叶黄素循环的非幅射能量耗散是银杏叶片防止强光损伤的主要机制     

光合作用光抑制的研究进展

概述了植物光合作用光抑制的研究进展,包括造成光抑制和光氧化的活性氧的产生和作用机理,光抑制的作用部位,以及光保护机制等,着重从三个方面讨论了植物抗光抑制的保护机理：与光系统Ⅱ天线以及叶黄素循环相关的热耗散途径,包括光呼吸、H2O-H2O循环和环式电子传递在内的电子传递途径,以及活性氧清除机制等。     

环境强光诱导玉簪叶片光抑制的机制

为进一步阐述光抑制的强光诱导和发生机制, 该文以喜阴植物玉簪(Hosta spp.)为材料研究其光抑制发生规律及其与环境光强的关系。结果表明, 全日照和遮阴条件下玉簪叶片发育分别形成适应强光和弱光的形态特征; 与遮阴处理相比, 强光下生长的玉簪光合速率和叶绿素含量较低, 但两种处理叶片最大光化学效率差异很小, 证明强光下植株可以正常生长且光合机构未发生严重的光抑制。将遮阴处生长的植株转移到全日照下, 光合速率和最大光化学效率急剧下降; 荧光诱导动力学曲线发生明显改变, 而且光系统II供体侧和受体侧荧光产量的变化幅度分别达到24.3%和34.2%, 表明玉簪由弱光转入强光后光系统II发生不可逆失活, 且受体侧受到的伤害较供体侧更严重。因此, 作者认为环境光强骤然提高并超过玉簪生长光强时很容易诱导其光合机构发生严重的光抑制。该研究对于理解植物适应光环境的策略以及喜阴植物的优质栽培有重要意义。     

低氧胁迫下黄瓜植株热耗散途径

采用营养液栽培,研究了低氧(营养液溶氧浓度为0.9～1.1 mg·L-1)胁迫下黄瓜幼苗光合作用热耗散与叶黄素循环的关系.结果表明:低氧胁迫下,黄瓜叶片PSⅡ的实际光化学效率(φPSⅡ)、饱和光强下的净光合速率(Pn)、表观量子效率(AQY)和PSⅡ的最大光化学效率(Fv/Fm)均显著降低,表明黄瓜植株的光合作用受到了光抑制;同时,光化学猝灭系数(qp)降低,而热耗散(NPQ)和天线耗散能量(D)的比值显著升高,说明黄瓜叶片热耗散增强;NPQ与叶黄素脱环氧化状态(DEPS)呈显著正相关,且两者均被抗坏血酸(AsA)所促进,被二硫苏糖醇(DTT)所抑制,说明低氧胁迫下,叶黄素循环是黄瓜植株光合作用热耗散的主要途径.     

光胁迫下银杏光合作用的光抑制

自然条件下晴天银杏叶片光系统Ⅱ光化学效率表现明显日变化。上午Ｆｖ／Ｆｍ随光照的增强而降低,至１４：００左右达最低值。其后随着光强的减弱Ｆｖ／Ｆｍ缓慢恢复。一天中叶黄素循环关键组分玉米黄质（Ｚ）含量与Ｆｖ／Ｆｍ呈负相关,用二硫苏糖醇（ＤＴＴ）阻断Ｚ的形成后,光抑制程度大大加深。结果表明与叶黄素循环有关的非辐射能量耗散的增加是产生光抑制的原因之一。强光处理前饲喂Ｄ１蛋白合成抑制剂林可霉素（ＬＭ）,ＦＶ     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

The photosynthesis of young Panicum C4 leaves is not C3-like

Evidence is presented contrary to the suggestion that C₄ plants grow larger at elevated CO₂ because the C₄ pathway of young C₄ leaves has C₃-like characteristics, making their photosynthesis O₂ sensitive and responsive to high CO₂. We combined PAM fluorescence with gas exchange measurements to examine the O₂ dependence of photosynthesis in young and mature leaves of Panicum antidotale (C₄, NADP-ME) and P. coloratum (C₄, NAD-ME), at an intercellular CO₂ concentration of 5 Pa. P. laxum (C₃) was used for comparison. The young C₄ leaves had CO₂ and light response curves typical of C₄ photosynthesis. When the O₂ concentration was gradually increased between 2 and 40%, CO₂ assimilation rates (A) of both mature and young C₄ leaves were little affected, while the ratio of the quantum yield of photosystem II to that of CO₂ assimilation (Φ_PSII/Φ_CO2) increased more in young (up to 31%) than mature (up to 10%) C₄ leaves. A of C₃ leaves decreased by 1·3 and Φ_PSII/Φ_CO2 increased by 9-fold, over the same range of O₂ concentrations. Larger increases in electron transport requirements in young, relative to mature, C₄ leaves at low CO₂ are indicative of greater O₂ sensitivity of photorespiration. Photosynthesis modelling showed that young C₄ leaves have lower bundle sheath CO₂ concentration, brought about by higher bundle sheath conductance relative to the activity of the C₄ and C₃ cycles and/or lower ratio of activities of the C₄ to C₃ cycles.     

Photosynthetic responses of Coffea arabica leaves to a short-term high light exposure in relation to N availability

It is known that the coffee (Coffea arabica L.) plant which is originally from shade habitats would have a limited ability to grow under full sun. Previous work has shown that nitrogen fertilisation can reduce the leaf damage when the plants are exposed to high light intensities during several days. In the present work we aimed to study the effects of the high irradiance during the first hours and evaluate the positive contribution of nitrogen fertilisation in the case of short-term exposure to strong light. Young plants (1.5–2 years old) grown in 1.5 kg of a mixed soil were supplemented with a nutrient solution containing 15 mM nitrogen in the form of NH₄NO₃, every 7 days (2N treatment), 15 days (1N treatment) and 45 days (0N treatment). Top mature leaves were exposed to a photosynthetic photon flux density of 1 500 μmol m^?2 s^?1 for a maximal period of 8 h, and changes in photosynthesis and pigment composition were monitored along the period of high light exposure. Photosynthetic capacity, leaf conductance to water vapour, electron transport capacity and maximum carboxylation activity, as well as some leaf fluorescence parameters (minimal fluorescence, photochemical efficiency of PSII and quantum yield of photosynthetic electron transport) were reduced by the stress, with a generally stronger impact observed in the 0N plants. The photochemical quenching was affected only in the 0N plants, while the non-photochemical quenching increased in 2N plants but decreased in the 0N ones. The results showed that 2N plants presented a better initial status of the photosynthetic parameters and of the content of photoprotective pigments. Those plants showed ability to trigger some protective mechanisms, as observed by the tendency to increase the xanthophyll pool content, specially in zeaxanthin and in non-photochemical quenching. Also, protein content presented a tendency to increase after 1.5 h, which was maintained until the end of the high light period. We conclude that nitrogen availability is a key factor in the acclimation process to high light.     

'Photoinhibition' of Heliconia under natural tropical conditions: the importance of leaf orientation for light interception and leaf temperature

The influence of irradiance on photosynthesis under natural conditions was studied in aseasonal Singapore using three Heliconia taxa: H. rostrata, H. psittacorum × H. spathocircinata cv. Golden Torch and H. psittacorum cv. Tay. When grown under full sunlight, all three heliconias exhibited reduced phatosynthetic capacities and lowered chlorophyll content per leaf area as compared with those grown under intermediate and deep shade. A marked decrease in the chlorophyll fluorescence F_v/Fm ratio and an increase in photochemical quenching (1- q_p) and non-photochemical quenching (q_N) were observed in upper leaves of plants grown under full sunlight. Increases in q_N suggest that ‘photoinhibition’ (decreases in F_v/F_m) in Heliconia grown under natural tropical conditions are probably due to photoprotective energy dissipation processes. The quantum yield, the maximum photosynthetic rate, F_v/F_m and the chlorophyll content of upper leaves were lower than those of lower leaves on the same plants grown under full sunlight. Similarly, lower values were obtained for the tip (sun) portion than for the base (shaded) portion of the leaves. The changes in F_v/F_m and in the levels of (1 –q_p) in leaves grown under intermediate and deep shade were negligible in plants during the course of day. However, there was a steep decrease in F_v/F_m and an increase in the levels of (1 –q_p), along with an increase in incident light in the sun leaves. The lowest F_v/F_m and the highest level of (1 –q_p) indicated minimum PSII efficiency at midday in full sun. These results indicate that, in Heliconia, the top leaves (particularly leaf tips) experienced sustained decreases in PSII efficiency upon exposure to full sunlight. Although all three taxa exhibited sustained decreases in photosynthetic capacity in full sunlight, the sun leaves of ‘Tay’ showed higher photosynthetic capacity than those of the other two taxa. This could be due, at least in part, to the vertical leaf angle and smaller lamina area. When the upright leaves of ‘Tay’ were constrained to a horizontal angle, they exhibited lower PSII efficiency (F_vIF_m ratio), while horizontal leaves of ‘Rostrata’ and ‘Golden Torch’ inclined lo near-vertical angles showed increased efficiency. Thus, an increase in leaf angle helps to achieve a reduction in the sustained decrease in PSII efficiency by decreasing the levels of incident sunlight and subsequently the leaf temperature.     

The inhibited xanthophyll cycle is responsible for the increase in sensitivity to low temperature photoinhibition in rice leaves fed with glutathione

Exposure of intact rice leaves to an irradiance of 1000 μmol m⁻² s⁻¹ at 6 °C for 2 h caused severe photoinhibition of Photosystem II. The rate and extent of photoinhbition were greatly exacerbated in leaves fed with 10 mM reduced glutathione (GSH) or 10 mM cysteine. Analyses of antioxidant enzyme activities as well as the application of protein synthesis inhibitors revealed that the increased sensitivity to photoinhibition following GSH feeding was not related to its effect on cellular antioxidant systems. On the other hand, feeding with GSH markedly suppressed the formation of zeaxanthin and antheraxanthin via the xanthophyll cycle and its associated nonradiative energy dissipation in leaves chilled in high light, suggesting that the stimulating effect of exogenous GSH on photoinhibition may be attributable to its action on the xanthophyll cycle. In vitro experiments using isolated thylakoids indicated that GSH is a weak inhibitor of violaxanthin deepoxidation. The possible implications of these results are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthesis and photoinhibition in leaves of chlorophyll b-less barley in relation to absorbed light

The response of photosynthesis to absorbed light by intact leaves of wild-type ( Hordeum vulgare L. cv. Gunilla) and chlorophyll b -less barley ( H. vulgare L. cv. Dornaria, chlorina-f2²⁸⁰⁰) was measured in a light integrating sphere. Up to the section where the light response curve bends most sharply the responses of the b -less and wild-type barley were similar but not identical. Average quantum yield and convexity for the mutant light response curves were 0.89 and 0.90, respectively, times those of the wild-type barley. The maximum quantum yield for PSII photochemistry was also 10% lower as indicated by fluorescence induction kinetics (F_v/F_m). Just above the region where the light curve bends most sharply, photosynthesis decreased with time in the mutant but not in the wild-type barley. This decrease was associated with a decrease in F_v/F_m indicating photoinhibition of PSII. This photoinhibition occurred in the same region of the light response curve where zeaxanthin formation occurs. Zeaxanthin formation occurred in both the chlorophyll b -less and wild-type leaves. However, the epoxidation state was lower in the mutant than in the wild-type barley. The results indicate that chlorophyll b -less mutants will have reduced photosynthetic production as a result of an increased sensitivity to photoinhibition and possibly a lowered quantum yield and convexity in the absence of photoinhibition.     

The xanthophyll cycle and sustained thermal energy dissipation activity in Vinca minor and Euonymus kiautschovicus in winter

The influence of low temperature on the operation of the xanthophyll cycle and energy dissipation activity, as ascertained through measurements of chlorophyll fluorescence, was examined in two broad-leaved evergreen species, Vinca minor L. and Euonymus kiautschovicus Loessner. In leaves examined under laboratory conditions, energy dissipation activity developed more slowly at lower leaf temperatures, but the final, steady-state level of such activity was greater at lower temperatures where the rate of energy utilization (through photosynthetic electron transport) was much lower. The rate at which energy dissipation activity increased was similar to that of the de-epoxidation of violaxanthin to antheraxanthin and zea-xanthin at different temperatures. However, leaves in the field examined prior to sunrise on mornings following cold days and nights exhibited a retention of antheraxanthin and zeaxanthin that was associated with sustained decreases in photosystem II efficiency. We therefore suggest that this phenomenon of ‘photoinhibition’ in response to light and cold temperatures during the winter results from sustained photoprotective thermal energy dissipation associated with the xanthophyll cycle. Such retention of the de-epoxidized components of the xanthophyll cycle responded to day-to-day changes in temperature, being greatest on the coldest mornings (when photoprotective energy dissipation might be most required) and less on warmer mornings when photosynthesis could presumably proceed at higher rates.