棉花幼苗叶片光合特性对低温胁迫及恢复处理的响应

为研究低温逆境及恢复处理下棉花幼苗光合特性响应机制, 丰富棉花苗期在不同胁迫水平下的抗寒机制及为应对自然条件下突发的低温冷害提供理论依据, 以‘新陆早33号’ (冷敏感)及‘中棉所50号’ (高耐寒)两个品种为试材, 采用人工模拟低温方法, 研究不同温度和处理时间下棉花幼苗光合气体交换参数、光能转化及传递的表现和恢复能力, 并通过测定胁迫解除后叶片的光合光响应曲线来分析叶片对光环境的适应能力。结果表明, 在较低强度低温逆境(15 ℃或24 h胁迫)中, 叶片净光合速率(P_n)、气孔导度(G_s)、气孔限制值、胞间CO₂浓度、最大光化学效率、光适应下最大光化学效率、实际光化学效率、相对电子传递速率变化幅度较小, 且胁迫解除后可恢复正常, 此时叶片光系统II (PSII)光反应中心受损可逆, P_n下降主要因气孔闭合引起。随着胁迫强度增加, 各测试指标变化显著, 且恢复表现较差, 此时叶片PSII光反应中心光能吸收、转化和电子传递受到严重阻碍, P_n下降原因由气孔限制因素转变为非气孔限制因素。低温胁迫导致叶片对光辐射的利用能力下降, 随着胁迫温度降低, 叶片最大净光合速率、表观量子效率及光饱和点快速下降, 光补偿点及暗呼吸速率则呈上升趋势。低温胁迫可导致棉花幼苗叶片对光环境适应能力降低, PSII反应中心对激发能捕获、活跃化学能转化及光合电子传递速率快速下降, CO₂固定能力降低, 最终导致光合能力下降。强耐寒品种则能在低温逆境中保持较高的光能转化、电子传递和弱光利用效率, 亦可通过减少暗呼吸消耗和调整G_s降低幅度和速度来保持较高的光合速率, 提高恢复能力, 增强植株抗逆性。     

热锻炼对甘蓝幼苗叶片激发能分配的影响

以喜温凉的蔬菜甘蓝为试材,研究了热锻炼与对照甘蓝幼苗叶片光合速率和叶绿素荧光参数对高温胁迫的响应．结果表明,叶片温度在25-35℃之间,热锻炼苗和对照苗叶片叶绿素可变荧光(Ｆv)、光化学猝灭(qP)、非光化学猝灭(qN)、PSⅡ化学效率(ФPSⅡ)没有明显的变化;当叶温高于35℃时。热锻炼苗的Fv、qP和中ФPSⅡ均明显高于对照,37℃时Fv、qP和ФPSⅡ分别比对照高53％、24％和86％;qN较对照低22％,尤其是与光抑制(光破坏)有关的qNs明显降低,以维持较高的高能态猝灭(qNf)耗散过剩激发能。保护PSⅡ反应中心不受破坏。减轻了光抑制,这与热锻炼幼苗叶片在高温下具有较高的光合能力是一致的。     

用体内叶绿素a荧光诱导动力学鉴定番茄的抗冷性

研究了冷害温度对具有不同抗冷性品种的番茄叶片的体内叶绿素a荧光诱导动力曲线的影响。实验结果指出,在低温处理(8℃,5℃,2℃下,暗中24小时)后,番茄叶片的体内叶绿素a荧光诱导动力学曲线有了明显的改变,Fv／Fo值、Rfd值降低了,光系统II原初光能转换效率和潜在的光合活力均受到抑制。我们在苗期和开花期得到的实验结果均表明,在番茄叶片的叶绿素a荧光诱导动力学曲线和这些荧光参数改变的程度与该品种的已知抗冷性之间呈现较好的相关性。我们认为,体内叶绿素a荧光诱导动力学方法是鉴定番茄抗冷性的一个快速、灵敏和可靠的方法,并可用于其他绿色植物的抗冷性鉴定中。     

紫茎泽兰茎和叶片色素及叶绿素荧光相关参数对不同温度处理的响应差异

通过人工模拟低温(12 ℃)、常温(25 ℃)、高温(35 ℃)生境,对紫茎泽兰茎和叶片色素(叶绿素a,b,类胡萝卜素,花青素)含量和组成、叶绿素荧光参数包括最大荧光效率Fv/Fm、光系统II效率ФpsⅡ、光化学淬灭系数qP、非光化学淬灭系数NPQ、热耗散速率HDR进行了动态测定.结果表明:在低温和高温胁迫处理过程中,茎和叶片的色素含量和组成随时间变化趋势基本一致,但茎的变化幅度明显低于叶片.与此类似,茎和叶片叶绿素荧光参数在不同温度处理过程中的变化趋势一致,但是茎各指标的变化幅度普遍小于相应叶片的变化幅度:低温下,茎的ФpsⅡ和ETR较对照最大降低44%,而叶片降低超过60%;高温下,茎的ФpsⅡ和ETR较对照下降16%～57%;而叶片则下降50%～80%.其产生原因在于:在温度胁迫条件下,叶片获取光能用于光化学过程的份额(qP)大幅下降,用于热耗散的份额(NPQ)大幅上升,茎的情况相反,所获取光能用于光化学电子传递的份额较常温下更多、用于热耗散的减少,这使得茎的耗散速率(HDR)升高的幅度显著低于叶片的升高幅度(p<0 05).综合3个温度的测定结果,茎的叶绿素含量相当于叶片的1/3～1/6,茎的叶绿素a/b较叶片低20%左右,但是光合电子传导速率ETR与叶片相当,这使得茎的光合色素利用效率ETR/Chl远高于叶片.叶片和茎叶绿素荧光参数在不同温度处理下变化趋势一致、但叶片的变化幅度远大于茎的这一响应差异,使得在适宜温度下紫茎泽兰叶片光合对整体光合贡献增大,而在温度胁迫条件下茎的光合贡献增大,这种策略使得这一植物在适宜生境下通过叶片光合、快速生长迅速占据生境,而在逆境条件下茎等非同化器官光合贡献增加,有利于其在逆境中的保存.     

生长温度对烟草叶片环式电子传递活性的影响（英文）

高等植物的光合机构在环境胁迫条件下非常容易产生光抑制,环式电子传递在光合机构的光保护中发挥着重要的作用。但是,生长温度对环式电子传递的影响并不清楚。本研究测定了在24/18℃和32/26℃条件下生长40天的烟草(K326)叶片的气体交换、叶绿素荧光和P700氧化还原态的光响应曲线。结果表明,烟草叶片在两种生长温度下的的光合能力、光化学淬灭、非光化学淬灭和通过光系统II的电子传递速率(ETR II)均没有差异。但是,在强光条件下,生长在24/18℃的叶片比生长在32/26℃的具有更高的通过光系统I的电子传递速率(ETR I)和ETR I/ETR II比值。短时间的强光处理后,生长在24/18℃的叶片具有较高的光系统II最大量子产额(Fv/Fm),表明环式电子传递活性的上调有助于缓解生长在24/18℃的叶片光系统II受到的光损伤。综上所述,环式电子传递活性的增强是植物适应较低生长温度的重要策略。     

长期冷适应对黑皮果蔗光合光能利用的影响

以拔地拉(Saccharum officinarum L.cv.Badila)幼苗为材料,研究了长期(28 d)冷适应处理(白天/夜间:12℃/8℃)对其光合光能利用的影响。结果表明:无论是常温(25℃)还是低温(12℃)测定,冷适应处理明显降低了蔗苗净光合速率(Pn),增加了暗呼吸速率(Rd)和电子传递量子效率和碳同化量子效率比值(ΦPSII/ΦCO2);常温下,冷适应处理降低了不同光强下Fv'/Fm'(和NPQ反向变化)随ΦPSII下降而下降的速率,同时增加了光化学淬灭q P随ΦPSII下降而下降的速率;而低温下,不同光强下Fv'/Fm'及q P随ΦPSII下降而下降的速率与对照没有显著差异;长期冷适应处理通过光化学途径和非光化学途径耗散了光合机构多余激发能,改善了其光合冷敏性,对果蔗抗冷栽培和抗低温育种有重要参考价值。     

高温胁迫下苋菜的叶绿素荧光特性

为了探明高温胁迫对苋菜(Amaranthus tricolor L.)光合过程的影响,用不同温度(25、30、35、40、45℃)处理苋菜植株1h后,随即测定了其叶绿素荧光动力学参数和快速光响应曲线特征参数的变化.结果表明:40℃以上高温胁迫下,苋菜叶片的光系统Ⅱ(PSⅡ)潜在光化学效率(Fv/Fo)、最大光化学效率(Fv/Fm)下降;最大荧光(Fm)、光合电子传递速率(ETR)、PSⅡ实际光化学效率(Yield)、光化学淬灭系数(qP)也均有所下降;而初始荧光(F.)和非光化学淬灭系数(NPQ)在40℃以上高温胁迫下显著上升.叶绿素荧光快速光响应曲线测定结果表明,初始斜率α、最大相对电子传递速率ETRmax和半饱和光强Ik在40℃以上高温胁迫下有所下降.研究表明,40℃以上高温胁迫对苋菜的光能的吸收、转换、光合电子传递和强光耐受能力等均有一定的影响.     

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。     

经济海藻红毛菜原位光合作用日变化

利用光合气体交换和叶绿素a荧光技术测定了原位沉水和干出条件下红毛菜光合作用的日变化,结果显示与沉水藻体相比,中午干出藻体光合速率、光系统II最大光化学效率(Fv/Fm)、实际光化学效率(ΦPSⅡ)、光化学淬灭系数(qP),光响应曲线初始斜率(ɑETR)和相对电子传递速率(rETR)值下降更明显。干出藻体重新入水后,其叶绿素a荧光参数在两个小时内可以完全恢复到沉水藻体的水平。红毛菜叶绿素a、类胡萝卜素、藻红蛋白的含量在一天中并没有出现明显变化。以上结果表明红毛菜日生长在中午要经历光抑制过程,干出状态下光抑制更严重;干出和沉水藻体光合速率都可以在傍晚得以恢复;红毛菜光系统II反应中心可以通过增加热耗散和降低光合电子传递速率等策略来应对光抑制。     

甲基紫精对水稻不同耐冷品种叶绿素荧光参数的影响

研究了不同浓度甲基紫精(MV)对水稻幼苗进行浸根处理,对不同耐冷品种常温和低温下叶绿素荧光参数的影响.结果发现,常温下,低浓度短时间的MV处理引起水稻原初光能转换效率(Fv/Fm)、光合电子传递量子效率(ΦPSⅡ)和光化学猝灭系数(qP)较对照增加,高浓度、长时间的MV处理则使各参数下降并低于对照,耐冷品种可以维持较高的非光化学猝灭系数(NPQ).低温下只有5 μmol/L甲基紫精(简称MV5)短时间处理时两品种的Fv/Fm增加,各处理引起ΦPSⅡ和qP下降,低浓度、短时间MV处理时NPQ增加.这说明轻度氧化胁迫可以刺激水稻的光合能力增加,但氧化胁迫会加剧水稻的低温伤害,耐冷品种也可以通过维持较高的线性电子传递速率和热耗散来避免这种过氧化伤害,表现出强的抗冷性.     

Effect of cold hardening on sensitivity of winter and spring wheat leaves to short-term photoinhibition and recovery of photosynthesis

Photoinhibition of photosynthesis and its recovery were studied in wheat (Triticum aestivum L.) leaves grown at nonhardening (20°C) and cold-hardening (5°C) temperatures. Cold-hardened wheat leaves were less susceptible to photoinhibition at 5°C than nonhardened leaves, and the winter cultivars, Kharkov and Monopol, were less susceptible than the spring cultivar, Glenlea. The presence of chloramphenicol, a chloroplastic protein synthesis inhibitor, increased the susceptibility to photoinhibition, but cold-hardened leaves still remained less susceptible to photoinhibition than nonhardened leaves. Recovery at 50 μmol m⁻² s⁻¹ photosynthetic photon flux density and 20°C was at least biphasic, with a fast and a slow phase in all cultivars. Cold-hardened leaves recovered maximum fluorescence and maximum variable fluorescence in the dark-adapted state during the fast phase at a rate of 42% h⁻¹ compared with 22% h⁻¹ for nonhardened leaves. The slow phase occurred at similar rates (2% h⁻¹) in cold-hardened and nonhardened leaves. Full recovery required up to 30 h. Fast-recovery phase was not reduced by either lowering the recovery temperature to 5°C or by the presence of chloramphenicol. Slow-recovery phase was inhibited by both treatments. Hence, the fast phase of recovery does not require de novo chloroplast protein synthesis. In addition, only approximately 60% of the photochemical efficiency lost through photoinhibition at 5°C was associated with lost [¹⁴C]atrazine binding and, hence, with damage to the secondary quinone electron acceptor for photosystem II-binding site. We conclude that the decrease in susceptibility to photoinhibition exhibited following cold hardening of winter and spring cultivars is not due to an increased capacity for repair of photoinhibitory damage at 5°C but reflects intrinsic properties of the cold-hardened photosynthetic apparatus. A model to account for the fast component of recovery is discussed.     

Photoinhibition and Photoprotection in Ginkgo biloba leaves: Influence of Temperature,CO2 and O2

In midday ginkgo ( Ginkgo biloba L. ) leaves have to bear photon flux density over 1 400 μmol·m-2·s-l in combination with high temperatures around 35℃ at natural habitat. They show typical midday depression of stomatal conductance and of CO2 assimilation rate. The zeaxanthin changes with light intensity during the day. The influence of the combination of strong light and temperature on photoinhibition was also examined in the laboratory. A low CO2 internal conductance (31 mmol· m- 2·s- 1 ) was found in ginkgo leaves, which had been exposed to excessive light at temperature between 15 ℃ and 35 ℃ with reduced CO2 (80 μL·L-l) or oxygen (2%) for 2 h, causing a low CO2 concentration at the carboxylation site and a high proportion of photorespimtion. The ratio of electron transport to CO2 fixation was rather high in ginkgo ( 16 e- /CO2 at 25 ℃ ) as compared with other plants. It increased with temperature also in 2% 02 which could not be explained solely as due to change of photorespimtion. The reduction of oxygen in 340 or 80 μL·L- 1 CO2 had no effect on the extent of photoinhibition at all temperatures, which indicated that eleetron flow caused by photorespiration in excess light was negligible in protective effect in ginkgo leaves. However, a decreased CO2 coneentration increased photoinhibition, especially at high temperature. It is concluded that the dissipation of excessive excitation energy in the PS II antennae through the xanthophyll cycle may be the major protective mechanism to preventing from the deteriorated effects of strong light in ginkgo leaves.     

Effects of low temperature acclimation upon photosynthetic induction in barley primary leaves

Oxygen evolution and chlorophyll fluorescence were measured in cold-hardened and unhardened leaves of barley ( Hordeum vulgare L. cv. Asa) during the induction period of photosynthesis. The lag phase of light-saturated photosynthesis was increased and steady-state rates of photosynthesis were higher in cold-hardened than in unhardened barley leaves. Fluorescence was quenched more rapidly during the first minutes of induction in hardened than unhardened leaves, largely because of greater energy-dependent quenching (q_E). Also, slow fluorescence transients through the M peak were delayed and less pronounced in cold-hardened than in unhardened leaves. Based upon the combined fluorescence and oxygen evolution data it was concluded that cold-hardening delayed light activation of the energy consuming carbon reduction cycle, thereby delaying the use of ATP and NADPH formed in the light reaction. Measurements of oxygen evolution and fluorescence kinetics during photosynthetic induction under oxygenic and anoxygenic conditions suggest that oxygen photoreduction is important for additional ATP generation during both the onset of photosynthetic carbon assimilation and during steady-state photosynthesis.     

Photoinhibition of isolated mesophyll cells from cold-hardened and non-hardened winter rye

Cold-hardened rye leaves have been shown to be more resistant to low temperature photoinhibition than non-hardened rye leaves. Isolated mesophyll cells from winter rye (Secale cereale L. cv. Musketeer) were exposed to photoinhibitory light conditions to estimate the importance of leaf morphology and leaf optical properties in the resistance of cold-hardened rye leaves to photoinhibition. Cold-hardened rye cells showed more resistance to photoinhibition than non-hardened rye cells when monitored with chlorophyll a variable to maximal fluorescence ratio (F_v/F_m). Thus, leaf morphology does not contribute to the resistance of cold-hardened rye leaves to low temperature photoinhibition. However, cold-hardened and non-hardened rye cells showed a similar extent of photoinhibition when photsynthetic CO₂ fixation rates were measured. They also showed the same capacity to recover from photoinhibition. During both photoinhibition and recovery, F_v/F_m and light limited CO₂ fixation rates showed different kinetics. We propose that inactivation and subsequent reactivation during recovery of some light activated Calvin cycle enzymes explain the greater extent of photoinhibition of light limited CO₂ fixation and its faster recovery compared to F_v/F_m kinetics during photoinhibition.     

Cold-hardening-induced resistance to photoinhibition of photosynthesis in winter rye is dependent upon an increased capacity for photosynthesis

Analyses of chlorophyll fluorescence and photosynthetic oxygen evolution were conducted to understand why cold-hardened winter rye (Secale cereale L.) is more resistant to photoinhibition of photosynthesis than is non-hardened winter rye. Under similar light and temperature conditions, leaves of cold-hardened rye were able to keep a larger fraction of the PS II reaction centres in an open configuration, i.e. a higher ratio of oxidized to reduced Q_A (the primary, stable quinone acceptor of PSII), than leaves of non-hardened rye. Three fold-higher photon fluence rates were required for cold-hardened leaves than for non-hardened leaves in order to establish the same proportion of oxidized to reduced Q_A. This ability of cold-hardened rye fully accounted for its higher resistance to photoinhibition; under similar redox states of q_a cold-hardened and non-hardened leaves of winter rye exhibited similar sensitivities to photoinhibition. Under given light and temperature conditions, it was the higher capacity for light-saturated photosynthesis in cold-hardened than in non-hardened leaves, which was responsible for maintaining a higher proportion of oxidized to reduced Q_A. This higher capacity for photosynthesis of cold-hardened leaves also explained the increased resistance of photosynthesis to photoinhibition upon cold-hardening.Abbreviations F_m and F'_m fluorescence when all PSII reaction centres are closed in dark- and light-acclimated leaves, respectively - F_o and F'_o fluorescence when all PSII reaction centres are open in darkness and steady-state light, respectively - F_v variable fluorescence (F'_m-F'_o) under steady-state light conditions - F_v/F_m the ratio of variable to maximum fluorescence as an expression of the maximum photochemical yield of PSII in dark-acclimated leaves - Q_A the primary, stable, quinone electron acceptor of PSII - q_N non-photochemical quenching of fluorescence due to high energy state (pH) - q_p photochemical quenching of fluorescence - RH cold-hardened rye - RNH non-hardened rye This work was supported by a Natural Sciences and Engineering Research Council of Canada (NSERCC) Operating Grant to N.P.A.H. G.Ö. was supported by an NSERCC International Exchange Award and by the Swedish Natural Science Research Council.     

Low-Temperature Effects on Photosynthesis and Correlation with Freezing Tolerance in Spring and Winter Cultivars of Wheat and Rye

Winter cultivars of rye (Secale cereale L., cv Musketeer) and wheat (Triticum aestivum L. cvs Kharkov and Monopol), but not a spring cultivar of wheat (Glenlea), grown at cold-hardening temperatures showed, at high irradiances, a higher proportion of oxidized to reduced primary, stable quinone receptor (QA) than did the same cultivars grown under nonhardening conditions. In addition, there was a positive correlation between the effects of low-growth temperature on this increased proportion of oxidized QA, and a concomitant increase in the capacity for photosynthesis, and LT50, the temperature at which 50% of the seedlings are killed, in cultivars showing different freezing tolerances. This suggests that low-temperature modulation of the photosynthetic apparatus may be an important factor during the induction of freezing resistance in cereals. Finally, the control of photosystem II photochemistry by nonphotochemical quenching of excitation energy was identical for nonhardened and cold-hardened winter rye. However, examination of measuring temperature effects per se revealed that, irrespective of growth temperature, nonphotochemical quenching exerted a stronger control on photosystem II photochemistry at 10[deg] C rather than at 20[deg] C.     

Mitochondria contribute to increased photosynthetic capacity of leaves of winter rye (Secale cereale L.) following cold-hardening

Cold-hardening of winter rye (Secale cereale L. cv. Musketeer) increased dark respiration from ?2.2 to ?3.9 μmol O₂ m^?2s^?1 and doubled light-and CO₂-saturated photosynthesis at 20°C from 18.1 to 37.0μmol O₂ m^?2 s^?1 We added oligomycin at a concentration that specifically inhibits oxidative phosphorylation to see whether the observed increase in dark respiration reflected an increase in respiration in the light, and whether this contributed to the enhanced photosynthesis of cold-hardened leaves. Oligomycin inhibited light- and CO₂-saturated rates of photosynthesis in non-hardened and cold-hardened leaves by 14 and 25%, respectively, and decreased photochemical quenching of chlorophyll a fluorescence to a greater degree in cold-hardened than in non-hardened leaves. These data indicate an increase both in the rate of respiration in the light, and in the importance of respiration to photosynthesis following cold-hardening. Analysis of metabolite pools indicated that oligomycin inhibited photosynthesis by limiting regeneration of ribulose-1,5-bisphosphate. This limitation was particularly severe in cold-hardened leaves, and the resulting low 3-phospho-glycerate pools led to a feed-forward inhibition of sucrose-phosphate synthase activity. Thus, it does not appear that oxidative phosphorylation supports the increase in photo-synthetic O₂ evolution following cold-hardening by increasing the availability of cytosolic ATP. The data instead support the hypothesis that the mitochondria function in the light by using the reducing equivalents generated by non-cyclic photosynthetic electron transport.     

基于叶绿素荧光成像及荧光参数分布特征的叶片光合异质性定量分析

植物叶片光合作用具有高度的空间异质性,叶绿素荧光成像技术为叶片光合异质性的研究提供了便利,但叶片光合异质性的定量分析并没有得到广泛应用。本文利用ImagingPAM叶绿素荧光成像系统,获得 中亚热带地区越冬期小叶榕（Ficus microcarpa）阳生叶和阴生叶的叶绿素荧光参数图像,并利用仪器的分析软件对其进行分析,定量比较了阳生叶和阴生叶的光合异质性特征。研究发现：越冬期小叶榕阳生叶的光合异质性和光抑制程度明显高于阴生叶,变异系数可作为光合异质性的定量指标。低温强光导致阳生叶坏死率（P_LN）达4.30%,并有53.30%的区域处于严重光抑制（0<F_v/F_m<0.627）,但仍有42.27%的区域仅为轻度光抑制（0.627≤ F_v/F_m<0.800）。而低温弱光并未造成阴生叶坏死和严重光抑制。通过对光系统Ⅱ（PSⅡ）的实际光合效率 （Y（Ⅱ））、调节性能量耗散的量子产额（Y（NPQ））和非调节性能量耗散的量子产额（Y（NO））荧光参数异质性的定量分析表明,阳生叶具有相对较高的光化学能力,阴生叶则具有相对较高的热耗散能力;冬季强光虽然会导致小叶榕阳生叶PSⅡ严重激发压积累,存在严重光抑制的潜在风险,但其致死面积并不大,叶片中仍存在一定面积低激发压的低风险区,而低温弱光下的阴生叶则主要以低风险区域为主。     

Photosynthetic acclimation to low temperature by western red cedar seedlings

Imposition of low, but above freezing, temperatures resulted in a gradual increase in the cold hardiness of western red cedar seedlings. This was associated with a decrease in the maximum rates of photosynthetic CO₂ fixation and O₂ evolution, and changes in chlorophyll a fluorescence transients which indicated that photoinhibition had occurred. Maximum photosynthetic rates declined approximately 40% during cold hardening. The leaves changed colour from green to red-brown during the hardening process. The colour change was due to the synthesis of large amounts of the carotenoid rhodoxanthin. Lutein levels doubled, while chlorophyll declined slightly. Dehardening resulted in the rapid recovery of photosynthesis to control levels, the rapid disappearance of rhodoxanthin, and the return of lutein levels to control. It is suggested that rhodoxanthin accumulation at low temperature functions to decrease the light intensity reaching the photosynthetic apparatus. The combination of photoinhibition and rhodoxanthin synthesis probably serves to protect the photosynthetic capacity of the seedlings at low temperature.     

Photoinhibition and photoprotection in senescent leaves of field-grown wheat plants

Photosynthesis, photosystem II (PSII) photochemistry, photoinhibition and the xanthophyll cycle in the senescent flag leaves of wheat (Triticum aestivum L.) plants grown in the field were investigated. Compared to the non-senescent leaves, photosynthetic capacity was significantly reduced in senescent flag leaves. The light intensity at which photosynthesis was saturated also declined significantly. The light response curves of PSII photochemistry indicate that a down-regulation of PSII photochemistry occurred in senescent leaves in particular at high light. The maximal efficiency of PSII photochemistry in senescent flag leaves decreased slightly when measured at predawn but substantially at midday, suggesting that PSII function was largely maintained and photoinhibition occurred in senescent leaves when exposed to high light. At midday, PSII efficiency, photochemical quenching and the efficiency of excitation capture by open PSII centers decreased considerably, while non-photochemical quenching increased significantly. Moreover, compared with the values at early morning, a greater decrease in CO₂ assimilation rate was observed at midday in senescent leaves than in control leaves. The levels of antheraxanthin and zeaxanthin via the de-epoxidation of violaxanthin increased in senescent flag leaves from predawn to midday. An increase in the xanthophyll cycle pigments relative to chlorophyll was observed in senescent flag leaves. The results suggest that the xanthophyll cycle was activated in senescent leaves due to the decrease in CO₂ assimilation capacity and the light intensity for saturation of photosynthesis and that the enhanced formation of antheraxanthin and zeaxanthin at high light may play an important role in the dissipation of excess light energy and help to protect photosynthetic apparatus from photodamage. Our results suggest that the well-known function of the xanthophyll cycle to safely dissipate excess excitation energy is also important for maintaining photosynthetic function during leaf senescence.