干旱和复水对入侵植物三裂叶蟛蜞菊叶片叶绿素荧光特性的影响

选取入侵植物三裂叶蟛蜞菊(Wedelia trilobata)及其本地近缘种蟛蜞菊(W. chinensis)为实验材料,比较干旱和复水后二者叶片的叶绿素荧光特性和抗氧化酶活性等生理指标的变化规律,探讨入侵种三裂叶蟛蜞菊对干旱的响应和生态适应性.结果发现,在自然干旱处理过程中,入侵种三裂叶蟛蜞菊与本地种蟛蜞菊相比土壤含水量下降较快,对它们叶片气孔形态的比较发现,干旱胁迫11d后三裂叶蟛蜞菊叶片气孔开度明显大于蟛蜞菊,这可能是导致其失水较快的原因之一.干旱胁迫11d后三裂叶蟛蜞菊的PSⅡ最大光能转化效率(Fv/Fm)降低了43.8%,而蟛蜞菊只降低了3.7%;同时,与蟛蜞菊相比,三裂叶蟛蜞菊的PSⅡ实际光化学量子产量(Yield)、表观光合电子传递速率(ETR)和光化学猝灭系数(qP)也表现出较大幅度的降低,说明三裂叶蟛蜞菊对干旱胁迫较敏感;但复水后,三裂叶蟛蜞菊能够较快地恢复到正常水平,且与本地种不存在显著差异.这主要是由于入侵种在遭受干旱胁迫时提高了对其过量激发能的热耗散能力以及超氧化物歧化酶(SOD)和抗坏血酸还原酶(APX)的活性,保护光合机构少受不可逆的损伤,使其在干旱胁迫解除后光合功能得以迅速恢复.研究结果初步表明三裂叶蟛蜞菊容易受到水分条件的限制,它向干旱地区扩散的可能性较小.     

亚低温与干旱胁迫对番茄叶片光合及荧光动力学特性的影响

利用人工气候室,研究了亚低温(8～15℃)和干旱胁迫(田间持水量的55％ ～ 65％)对盆栽番茄叶片光合特性、能量及电子流分配的影响.结果表明:与对照相比,亚低温胁迫下番茄叶片的光合色素含量降低,而干旱胁迫下升高.在亚低温或干旱胁迫下,番茄叶片的胞间CO2浓度、气孔导度及净光合速率均显著下降,气孔限制值升高,其共同处理下相应指标进一步降低或升高.亚低温或干旱单一胁迫提高了光呼吸速率,但共同胁迫下其光呼吸速率反而降低.无论是亚低温、干旱胁迫还是共同胁迫,均会导致初始荧光、PSⅡ原初光能转化效率和有效光量子产量下降,光系统发生损伤,并且PSⅡ激发能分配系数升高,光化学效率降低,过剩光能增加,总电子流及流向各交替电子流库的电子流减少.为耗散过剩光能,热耗散及交替电子流的比例增加.与亚低温和干旱胁迫单一处理相比,二者共同处理加剧了叶片的热耗散,但交替电子流比例没有进一步增加.     

Effects of water stress on photochemical function and protein metabolism of photosystem II in wheat leaves

The effects of osmotic dehydration in wheat leaves ( Triticum aestivum L. cv. Longchun No. 10) on the photochemical function and protein metabolism of PSII were studied with isolated thylakoid and PSII membranes. The results indicated that PSII was rather resistant to water stress as mild water deficit in leaves did nut significantly affect its activity. However, extreme stress conditions such as 40% decrease in relative water content (RWC) or 1.8 MPa in water potential (Ψ) caused ca 50% reduction in O₂ evolution and ca 25% inhibition of DCIP (2.6-dichlorophenol indophenol) photoreduction of PSII. In addition, it was found that the inhibited DCIP photoreduction of PSII could not be reversed by DPC (2.2-diphenylcarbazide), a typical electron donor to PSII, suggesting that water stress did not affect electron donation to PSII. Urea-SDS-PAGE and western blot analysis showed that the steady slate levels of major PSII proteins, including the D1 and D2 proteins in the PSII reaction center, declined on a chlorophyll basis with increasing water stress, possibly as a result of increased degradation. In vitro translation experiments and quantitative analysis of chloroplast RNAs indicated that the potential synthesis of chloroplast proteins from their mRNAs was impaired by water stress. From the results it is concluded that the effects of water stress on PSII protein metabolism, especially on the reaction center proteins, may account for the damage to PSII photochemistry.     

Effects of water stress on major photosystem II gene expression and protein metabolism in barley leaves

Although it has long been recognized that water deficit in plants reduces photosystem (PS) II mRNAs and proteins, the detailed mechanisms behind this have not been thoroughly elucidated. In the present study, effects of water stress in barley leaves on degradation of major PSII mRNA and dissociation and migration of PSII proteins were investigated. The results indicated that (1) the steady-state levels of major PSII mRNAs and proteins declined with increasing water stress, as a consequence of increased degradation; under severe water stress, the half-lives of D1 and D2 proteins decreased from 12–14 h to 7–8 h and the half-lives of psbA and psbD mRNA decreased from above 16 to 6–10 h; (2) monomerization of PSII were increased during water stress. Severe water stress accelerated turnover of PSII and inhibited PSII activities.     

氮调控对盐环境下甜菜功能叶光系统Ⅱ荧光特性的影响

采用盆栽试验方法,以NaCl为盐分模拟不同盐度环境,研究了施氮(N)对盐环境下生长的甜菜(Beta vulgaris)功能叶光系统Ⅱ (PSⅡ)荧光特性的影响及光合色素含量的变化.结果表明:在轻度、中度及重度盐环境下,施N均能增大PSⅡ最大光化学效率(Fv/Fm)、PSⅡ潜在活性(Fv/Fo)、PSⅡ实际光量子产量(Y(Ⅱ))、非调节性能量耗散的量子产量(Y(NO))、相对电子传递速率(ETR)及光化学猝灭系数(qp),且在适宜的施N范围内(0-1.2 g·kg-1)上述参数随施N量的增加而增大.各叶绿素荧光参数光响应的结果表明,随着光强的增加,各处理下调节性能量耗散的量子产量(KNPQ))、ETR及非光化学猝灭系数(NPQ)旱上升趋势,相反,Y(Ⅱ)、Y(NO)及qp则呈下降趋势,在有效的光强范围内(0-1 000 μmol·m-2·s-1)施N提高了甜菜功能叶PSⅡ反应中心的开放程度,并且在高光强下调节PSⅡ耗散掉过剩的光能以避免对其反应中心造成伤害.各盐度环境下施N也显著增加了甜菜功能叶叶绿素与类胡萝卜素含量,增大了叶绿素a/叶绿素b值,且叶绿素与类胡萝卜素含量随施N水平的增加而增加.说明盐环境下施N能够增强甜菜功能叶PSⅡ的活性,提高PSⅡ光能利用率,从而增强其对盐渍环境的适应性.     

Drought stress effects on photosystem I content and photosystem II thermotolerance analyzed using Chl a fluorescence kinetics in barley varieties differing in their drought tolerance

Drought stress has multiple effects on the photosynthetic system. Here, we show that a decrease of the relative contribution of the I-P phase, ΔV_IP = − V _I = ( F _M− F _I)/(F_M− F_o), to the fluorescence transient OJIP is observed in 10 drought-stressed barley and 9 chickpea varieties. The extent of the I-P loss in the barley varieties depended on their drought tolerance. The relative loss of the I-P phase seems to be related to a loss of photosystem (PS) I reaction centers as determined by 820-nm transmission measurements. In the second part of this study, the interaction of drought and heat stress in two barley varieties (the drought tolerant variety A¨t Baha and the drought sensitive variety Lannaceur) was studied using a new approach. Heat stress was induced by exposing the plant leaves to temperatures of 25–45°C and the inactivation of the O₂-evolving complex (OEC) was followed measuring chlorophyll a (Chl a ) fluorescence using a protocol consisting of two 5-ms pulses spaced 2.3 ms apart. In active reaction centers, the dark interval is long enough to allow the OEC to recover from the first pulse; whereas in heat-inactivated reaction centers it is not. In the latter category of reaction centers, no further fluorescence rise is induced by the second pulse. Lannaceur, under well-watered conditions, was more heat tolerant than Aït Baha. However, this difference was lost following drought stress. Drought stress considerably increased the thermostability of PS II of both varieties.     

Long-term drought stress induces structural and functional reorganization of photosystem II

Long-term drought stress on photosystem II (PSII) was studied in pea (Pisum sativum L.) seedlings. Drought stress (reduction of water content by 35–80%) led to a considerable depletion of the PSII core, and the remaining PSII complex appeared to be functional and reorganized, with a unit size (LHCP/PSII core) twofold greater than that of well-irrigated plants. By immunoblotting analysis of the PSII proteins from grana and stroma lamellae, the enhanced degradation of CP43 and D1 proteins was observed in water-stressed plants. Also, water stress caused increased phosphorylation of the PSII core and increased D1 protein synthesis. Water-stress-mediated increase in D1 synthesis did not occur when plants were exposed to photoinhibitory light. The depletion of the PSII core was essentially reversed when water-stressed plants grown at low visible irradiance were watered. We suggest that the syndrome caused by the effect of long-term water stress on photosynthesis is a combination of at least two events: a reduction in the number of active PSII centres caused by a physical destabilization of the PSII core and a PSII reorganization with enhanced D1 turnover to counteract the core depletion.Abbreviations Chl chlorophyll - CP43 and CP47 -carotene-Chla-proteins of PSII core - DCPIP 2,6-dichlorophenolindophenol - DPC diphenylcarbazide - Fv/Fm the ratio of yield of variable fluorescence to yield of maximal fluorescence when all reaction centres are closed - LHC(P) light-harvesting complex (proteins) - Wc water content This work was supported by the Italian National Council of Research special grant RAISA, subproject 2 (paper No. 2179) on water stress B. Geiken was supported by the European program Human Capital and Mobility. We thank Dr. Roberto Barbato (Department of Biology, University of Padua, Italy) for generous gifts of various PSII antibodies.     

NaCl胁迫加重强光胁迫下超大甜椒叶片的光系统II和光系统I的光抑制

快速叶绿素荧光动力学可以在无损情况下探知叶片光合机构的损伤程度, 快速叶绿素荧光测定和分析技术(JIP-test)将测量值转化为多种具有生物学意义的参数, 因而被广泛应用于植物光合机构对环境的响应机制研究。该文研究了超大甜椒(Capsicum annuum)幼苗在强光及不同NaCl浓度胁迫下的荧光响应情况。与单纯强光胁迫相比, NaCl胁迫引起了叶绿素荧光诱导曲线的明显改变, 光系统II (PSII)光抑制加重, 同时PSII反应中心和受体侧受到明显影响, 而且高NaCl浓度胁迫下PSII供体侧受伤害明显, 同时PSI反应中心活性(P700⁺)在盐胁迫下明显降低。这些结果表明, NaCl胁迫会增强强光对超大甜椒光系统的光抑制, 并且浓度越高抑制越明显, 但对PSI的抑制作用低于PSII。高NaCl浓度胁迫易对PSII供体侧造成破坏, 且PSI光抑制严重。     

外源ATP对NaCl胁迫下菜豆叶片叶绿素荧光特性的调节

盐胁迫是影响植物生长的主要逆境因子之一,外源ATP被发现可作为信号分子参与植物对逆境胁迫生理反应的调节。为了探明外源ATP在植物盐胁迫响应中的作用,以增强植物对土壤盐渍化的耐性,更好地应用于土壤盐渍化修复。该研究以菜豆( Phaseolus vulgaris)为材料,通过叶绿素荧光技术探讨了外源ATP 对菜豆叶片在NaCl胁迫下叶绿素荧光特性的变化规律。结果表明：在NaCl胁迫下,叶片光系统Ⅱ( PSⅡ)潜在最大光化学量子效率( Fv/Fm)、光适应下最大光化学效率( Fv′/Fm′)、PSⅡ光适应下实际光化学效率[ Y (Ⅱ)]、光化学荧光猝灭( qP)、电子传递速率( ETR)与对照组相比均有显著性下降,而非光化学猝灭( NPQ)和( qN)较对照组有显著性增加,这表明NaCl胁迫导致菜豆叶片光系统Ⅱ光化学效率的下降和光能耗散的增加。而外源ATP(eATP)的处理能有效缓解NaCl胁迫所造成的Fv/Fm、Fv′/Fm′、Y(Ⅱ)、qP、ETR下降和NPQ、qN的上升。该研究结果表明在NaCl胁迫下外源ATP可以有效地提高菜豆幼苗光系统Ⅱ( PSⅡ)的光化学反应效率。     

Time- and reduction-dependent rise of photosystem II fluorescence during microseconds-long inductions in leaves

Photosynthesis Research - Lettuce (Lactuca sativa) and benth (Nicotiana benthamiana) leaves were illuminated with 720 nm background light to mix S-states and oxidize electron carriers....     

Effects of heat and high irradiance stress on energy dissipation of photosystem II in low irradiance-adapted peanut leaves

To increase crop yields and not to compete for land with food crops, intercropping agricultural cultivation approach was introduced into cultivation of peanut (Arachis hypogaca L.). This approach improves the total yield of the crop per unit area, but decreases the yield of a single crop compared with mono-cropped agricultural cultivation approach. In wheat-peanut relay intercropping system, peanut plants would suffer heat and high light (HI) stress after wheat harvest. In the present work, peanut seedlings were cultivated in low light to simulate wheat-peanut relay intercropping environments. Upon exposure to heat and HI stress, energy dissipation in PSII complexes was evaluated by comparing those cultivated in low irradiance conditions with the mono-cropped peanut. The maximal photochemical efficiency of PSII (F_v/F_m) and the net photosynthetic rate (P_n) decreased markedly in relay-cropped peanut (RP) after heat and HI stress, accompanied by higher degree of PSII reaction center closure (1–qP). After heat and HI stress, higher antioxidant enzyme activity and less ROS accumulation were observed in mono-cropped peanut (MP) seedlings. Meanwhile, higher content of D1 protein and higher ratio of (A + Z)/(V + A + Z) were also detected in MP plants under such stress. These results implied that heat and HI stress could induce photoinhibition of PSII reaction centers in peanut seedlings and both xanthophyll cycle-dependent thermal energy dissipation and the antioxidant system were down-regulated in RP compared to classical monocropping systems after heat and high irradiance stress.     

Evaluation of chlorophyll fluorescence as a probe for drought stress in willow leaves

The effect of drought on the photosynthetic apparatus of leaves of Salix sp. was studied by measurements of the induction of chlorophyll fluorescence and the capacity for O₂ evolution. Using a multivariate analysis, a model was developed that could predict the degree of drought stress from the data of fluorescence kinetics. Even mild drought stress was detected with high precision; this was not always possible when the photosynthetic capacity was measured. The most clear discrimination between control and drought-stressed leaves was obtained if fluorescence induction was measured at high rather than normal CO₂ levels, and at low rather than high light levels. All information provided by fluorescence pertaining to drought was contained within the slow phase of the induction curve. It is suggested that rapid dehydration is different from drought at the mechanistic level as judged by the fluorescence characteristics.     

外源钙对干旱胁迫下烤烟幼苗光系统Ⅱ功能的影响

以叶绿素快相荧光动力学曲线(OJIP)为探针,研究了外源钙对干旱胁迫下烤烟幼苗光系统Ⅱ(PSⅡ)功能的影响.结果表明:干旱胁迫降低了烤烟幼苗PSⅡ原初光能转换效率(Fv/Fm)和电子传递速率(ETR),抑制了光合作用的原初过程,烤烟幼苗叶片发生了明显的光抑制.叶面喷施10.0 mmol·L-1CaCl2溶液后烤烟叶片的光合电子传递能量比例(ФEo)在干旱胁迫下的降低幅度明显小于对照(喷施清水),电子转运效率(ET0/RC)在干旱胁迫下明显高于对照.叶面喷施CaC12溶液增加了PSⅡ捕获光能用于光合电子传递的比例、剩余有活性反应中心的效率和电子传递链中的能量传递,使烤烟叶片的光系统Ⅱ在干旱胁迫下保持相对较高的活性,从而提高了烤烟幼苗的抗旱能力.     

Responses of photosystem I compared with photosystem II to high-light stress in tropical shade and sun leaves

Sun and shade leaves of several plant species from a neotropical forest were exposed to excessive light to evaluate the responses of photosystem I in comparison to those of photosystem II. Potential photosystem I activity was determined by means of the maximum P700 absorbance change around 810 nm (ΔA_810max) in saturating far-red light. Leaf absorbance changes in dependence of increasing far-red light fluence rates were used to calculate a ‘saturation constant’, K_s, representing the far-red irradiance at which half of the maximal absorbance change (ΔA_810max/2) was reached in the steady state. Photosystem II efficiency was assessed by measuring the ratio of variable to maximum chlorophyll fluorescence, F_v/F_m, in dark-adapted leaf samples. Strong illumination caused a high degree of photo-inhibition of photosystem II in all leaves, particularly in shade leaves. Exposure to 1800–2000 μ mol photons m⁻² s⁻¹ for 75 min did not substantially affect the potential activity of photosystem I in all species tested, but caused a more than 40-fold increase of K_s in shade leaves, and a three-fold increase of K_s in sun leaves. The increase in K_s was reversible during recovery under low light, and the recovery process was much faster in sun than in shade leaves. The novel effect of high-light stress on the light saturation of P700 oxidation described here may represent a complex reversible mechanism within photosystem I that regulates light-energy dissipation and thus protects photosystem I from photo-oxidative damage. Moreover, we show that under high-light stress a high proportion of P700 accumulates in the oxidized state, P700⁺. Presumably, conversion of excitation energy to heat by this cation radical may efficiently contribute to photoprotection.     

A model of photosystem II for the analysis of fast fluorescence rise in plant leaves

The polyphasic patterns of fluorescence induction rise in pea leaves in vivo and after the treatment with ionophores have been studied using a Plant Efficiency Analyzer. To analyze in detail photosystem II (PS II) electron transfer processes, an extended PS II model was applied, which included the sums of exponential functions to specify explicitly the light-driven formation of the transmembrane electric potential (ΔΨ(t)) as well as pH in the lumen (pH_L(t)) and stroma (pH_S(t)). PS II model parameters and numerical coefficients in ΔΨ(t), pH_L(t), and pH_S(t) were evaluated to fit fluorescence induction data for different experimental conditions: leaf in vivo or after ionophore treatment at low or high light intensity. The model imitated changes in the pattern of fluorescence induction rise due to the elimination of transmembrane potential in the presence of ionophores, when ΔΨ = 0 and pH_L(t), pH_S(t) changed to small extent relative to control values in vivo, with maximum ΔΨ(t) ∼ 90 mV and ΔΨ(t) ∼ 40 mV for the stationary state at ΔpH ≅ 1.8. As the light intensity was increased from 300 to 1200 μmol m⁻² s⁻¹, the heat dissipation rate constants increased threefold for nonradiative recombination of P680⁺Phe⁻ and by ∼30% for P680⁺Q_A⁻. The parameters ΔΨ, pH_S and pH_L were analyzed as factors of PS II redox state populations and fluorescence yield. The kinetic mechanism of fluorescence quenching is discussed, which is related with light-induced lumen acidification, when ⁺Q_A⁻ and P680⁺ recombination probability increases to regulate the Q_A reduction.     

Mechanism of strong quenching of photosystem II chlorophyll fluorescence under drought stress in a lichen, Physciella melanchla, studied by subpicosecond fluorescence spectroscopy

The mechanism of the severe quenching of chlorophyll (Chl) fluorescence under drought stress was studied in a lichen Physciella melanchla, which contains a photobiont green alga, Trebouxia sp., using a streak camera and a reflection-mode fluorescence up-conversion system. We detected a large 0.31 ps rise of fluorescence at 715 and 740 nm in the dry lichen suggesting the rapid energy influx to the 715-740 nm bands from the shorter-wavelength Chls with a small contribution from the internal conversion from Soret bands. The fluorescence, then, decayed with time constants of 23 and 112 ps, suggesting the rapid dissipation into heat through the quencher. The result confirms the accelerated 40 ps decay of fluorescence reported in another lichen (Veerman et al., 2007 [36]) and gives a direct evidence for the rapid energy transfer from bulk Chls to the longer-wavelength quencher. We simulated the entire PS II fluorescence kinetics by a global analysis and estimated the 20.2 ns^− 1 or 55.0 ns^− 1 energy transfer rate to the quencher that is connected either to the LHC II or to the PS II core antenna. The strong quenching with the 3-12 times higher rate compared to the reported NPQ rate, suggests the operation of a new type of quenching, such as the extreme case of Chl-aggregation in LHCII or a new type of quenching in PS II core antenna in dry lichens.     

Effects of drought on photosynthesis,chlorophyll fluorescence and photoinhibition susceptibility in intact willow leaves

Plants from clonal cuttings of Salix sp. were subjected to a drying cycle of 10 d in a controlled environment. Gas exchange and fluorescence emission were measured on attached leaves. The light-saturated photosynthetic CO₂ uptake became progressively inhibited with decreased leaf water potential both at high, and especially, at low intercellular CO₂ pressure. The maximal quantum yield of CO₂ uptake was more resistant. The inhibition of light-saturated CO₂ uptake at leaf water potentials around-10 bar, measured at a natural ambient CO₂ concentration, was equally attributable to stomatal and non-stomatal factors, but the further inhibition below this water-stress level was caused solely by non-stomatal factors. The kinetics of fluorescence emission was changed at severe water stress; the slow secondary oscillations of the induction curve were attenuated, and this probably indicates perturbations in the carbon reduction cycle. The influence of light level during the drought period was also studied. Provided the leaves were properly light-acclimated, drought at high and low light levels produced essentially the same effects on photosynthesis. However, low-light-acclimated leaves became more susceptible to photoinhibitory treatment under severe water stress, as compared with well-watered conditions.     

A model of photosystem II for the analysis of a fast fluorescence rise in plant leaves

The polyphasic patterns of fluorescence induction rise in pea leaves in vivo and after the treatment with ionophores have been studied using a plant efficiency analyzer. To analyze in detail photosystem II (PS II) electron transfer processes, an extended PS II model was applied, which included the sums of exponential functions to specify explicitly the light-driven formation of the transmembrane electric potential (delta psi(t)) as well as pH in the lumen (pHL(t)) and stroma (pHs(t)). PS II model parameters and numerical coefficients in delta psi(t), and pHs(t) were evaluated to fit fluorescence induction data for different experimental conditions: leaf in vivo or after ionophore treatment at low or high light intensity. The model imitated changes in the pattern of fluorescence induction rise due to the elimination of transmembrane potential in the presence of ionophores, when delta psi = 0 and pHL(t), pHS(t) altered to small extent relative to control values in vivo, with maximum delta psi(t) approximately 90 MB and delta psi(t) approximately 40 MB, for the stationary state at deltapH aproximately equal to 1.8. As the light intensity was increased from 300 to 1200 micromol x m(-2) x s(-1), the heat dissipation rate constants increased threefold for nonradiative recombination of P680+Phe- and by approximately 30% for P680+Q(A)-. The parameters delta psi, pH(S) and pH(L) were analyzed as factors of PS II redox state populations and fluorescence yield. The kinetic mechanism of qE quenching is discussed, which is related with light induced pH(L) lumen acidification, when Q(A)- and P680+ recombination probability increases to regulate the QA reduction.     

Effects of salinity stress on photosystem II function in cyanobacterial Spirulina platensis cells

The changes in PSII photochemistry in Spirulina platensis cells exposed to salinity stress (0–0.8 M NaCl) for 12 h were studied. Salinity stress induced a decrease in oxygen evolution activity, which correlated with the decrease in the quantum yield of PSII electron transport ( Φ _PSII). Phycocyanin content decreased significantly while chlorophyll content remained unchanged in salt-stressed cells. Salinity stress induced an increase in non-photochemical quenching (q_N) and a decrease in photochemical quenching (q_P). Analyses of the polyphasic fluorescence transients (OJIP) showed that with the increase in salt concentration, the fluorescence yield at the phases J, I and P declined sharply and the transient almost levelled off at salt concentration of 0.8 M NaCl. The effects of DCMU on the polyphasic rise of fluorescence transients decreased significantly. Salinity stress resulted in a decrease in the efficiency of electron transfer from Q_A⁻ to Q_B. The slope at the origin of the relative variable fluorescence curves (dV/dt_o) and the relative variable fluorescence at phase J (V_J) increased in the absence of DCMU, but decreased in the presence of DCMU. The shape of the relative variable fluorescence transients in salt-stressed cells was comparable to that of the control cells incubated with DCMU. The results in this study suggest that salt stress inhibited the electron transport at both donor and acceptor sides of PSII, resulted in damage to phycobilisome and shifted the distribution of excitation energy in favour of PSI.