Interaction of proline, sugars, and anthocyanins during photosynthetic acclimation of Arabidopsis thaliana to drought stress

The relationships among photosynthetic acclimation, proline (Pro), soluble sugar (SS), and anthocyanin (An) accumulation in Arabidopsis thaliana leaves to the onset of drought stress (OnDS), mild (MiDS) and moderate drought stress (MoDS), were evaluated. As leaf water content (LWC) decreased, metabolic concentrations (Pro, SS, and An) increased and were negatively and significantly correlated with LWC. Thus, these metabolites may have an important role in the acclimation process to drought stress (DS). No correlations among Pro, SS and An accumulation with the quantum efficiency of PSII photochemistry (Φ(PSII)) and the excitation pressure (1-q(P)) were observed under DS. This implies that, while metabolites increased in a drought-dependent way, PSII activity did not decrease in the same pattern. Our results indicated that, under MoDS, A. thaliana leaves were able to maintain oxidative compounds such as malondialdeyde, an end product of lipid peroxidation, within the range of control leaves, and to cope with oxidative damage, as was evident by the decreased excitation pressure (1-q(P)) and similar (ns difference) Φ(PSII) to that of control leaves. In addition, a statistically significant increased accumulation of Pro, SS and An was recorded only under MoDS compared to controls. The better PSII functioning of MoDS Arabidopsis leaves may reflect the greater capacity of these leaves to undertake key metabolic adjustments, including increased Pro, SS and An accumulation, to maintain a higher antioxidant protection and a better balance between light capture and energy use.     

Interactive Effects of Drought Stresses and Elevated CO2 Concentration on Photochemistry Efficiency of Cucumber Seedlings

To reveal and quantify the interactive effects of drought stresses and elevated CO2 concentration [CO2] on photochemistry efficiency of cucumber seedlings, the portable chlorophyll meter was used to measure the chlorophyll content, and the Imaging-PAM was used to image the chlorophyll fluorescence parameters and rapid light response curves (RLC) of leaves in two adjacent greenhouses. The results showed that chlorophyll content of leaves was reduced significantly with drought stress aggravated. Minimal fluorescence (Fo) was increased while maximal quantum yield of PSII (Fv/Fm) decreased significantly by severe drought stress. The significant decrease of effective quantum yield of PSll (Y(Ⅱ)) accompanied by the significant increase of quantum yield of regulated energy dissipation (Y(NPQ)) was observed under severe drought stress condition, but there was no change of quantum yield of nonregulated energy dissipation (Y(NO)). We detected that the coefficient of photochemical quenching (Qp) decreased, and non-photochemical quenching (NPQ) increased significantly under severe drought stress. Furthermore, we found that maximum apparent electron transport rate (ETRmax) and saturating photosynthetically active radiation (PPFDsat) decreased significantly with drought stress aggravated. However, elevated [CO2] significantly increased FvlFm, Qp and PPFDsat, and decreased NPQ under all water conditions, although there were no significant effects on chlorophyll content, Fo, Y(Ⅱ), Y(NPQ), Y(NO) and ETRmax. Therefore, it is concluded that CO2-fertilized greenhouses or elevated atmospheric [CO2] in the future could be favorable for cucumber growth and development, and beneficial to alleviate the negative effects of drought stresses to a certain extent.     

Spatial-temporal variations in rose leaves under water stress conditions studied by chlorophyll fluorescence imaging.

Spatial-temporal changes were examined by imaging chlorophyll (Chl) a fluorescence in four leaf areas, two central and two external of rose plants (Rosa x hybrida) cv. Grand Gala for 9 days, under progressive water stress. New fluorescence parameters based on the lake model have recently been used to determine Q(A) redox state and excitation energy fluxes in order to gain a better understanding of the mechanisms that occur under drought stress. Chlorophyll fluorescence images showed a spatial variation in the leaves. The lower values for F(o), F(M), phi(2), q(P) and q(L) were found in the internal leaf area while higher values of non-photochemical quenching calculated from Stern-Volmer quenching (NPQ) and phi(NPQ). phi(Po) were more homogeneous throughout leaf. Temporal changes were also observed during the experiment, a 10% decrease in relative water content (RWC) (between day 1 and 2), led to a decrease in photochemical quenching and an increase in non-photochemical processes. Chlorophyll fluorescence parameters were more or less constant till day 8. At the end of the experiment (day 9), energy dissipation by downregulation, electron transport and Q(A) redox state, decreased and phi(NO) increased to compensate the change. Chlorophyll fluorescence parameters based on the lake model q(L), phi(NPQ) and phi(NO) have been found more appropriate for estimating the fraction of open centres, the quantum yield of regulated energy dissipation in photosystem II (PSII) and the quantum yield of non-regulated energy dissipation in PSII, respectively. The F(s)/F(o) ratio is strongly correlated with NPQ and phi(NPQ) up to a RWC of 20%. This coincides with a greater decrease in photochemical quenching and non-photochemical quenching and an increase in phi(NO).     

塔里木河下游地下水位对柽柳叶绿素荧光特性的影响

选取塔里木河下游3处地下水埋深6m的监测井位作为研究点,结合典型生态监测断面的地下水位监测数据,分析不同地下水埋深处柽柳的叶绿素荧光特性和光系统的光合活性.结果表明:随着地下水埋深加大和干旱胁迫加剧,柽柳叶片的实际光化学效率、电子传输速率和光化学猝灭等参数普遍下降;非光化学猝灭和调节性能量耗散量子产量等参数显著升高,而最大光量子产量总体处于相对适宜状态.干旱胁迫下柽柳的PSII光合活性随地下水埋深增大而下降,捕获光能的过剩程度加剧,发生光抑制的几率增大,其自身良好的抗旱性和自我调节机制,使光系统II尚未发生显著光损伤.     

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.     

Operation of the xanthophyll cycle and degradation of D1 protein in the inducible CAM plant,Talinum triangulare,under water deficit

Changes in photochemical activity induced by water deficit were investigated in Talinum triangulare, an inducible CAM plant. The aim was to analyse the interactions between C3 photosynthesis, induction and activity of CAM, photosynthetic energy regulation and the mechanisms responsible for photoprotection and photoinhibition under water stress. Gas exchange, chlorophyll a fluorescence, titratable acidity, carotenoid composition and relative contents of the PSII reaction centre protein (D1) were measured. A decrease in xylem tension (psi) from -0.14 to -0.2 MPa substantially decreased daytime net CO2 assimilation and daily carbon gain, and induced CAM, as shown by CO2 assimilation during the night and changes in titratable acidity; a further decrease in psi decreased nocturnal acid accumulation by 60%. Non-photochemical quenching of chlorophyll a fluorescence (NPQ) increased with water deficit, but decreased with a more severe drought (psi below -0.2 MPa), when CAM activity was low. NPQ was lower at 0900 h (during maximum decarboxylation rates) than at 1400 h, when malate pools were depleted. Down-regulation of PSII activity related to the rise in NPQ was indicated by a smaller quantum yield of PSII photochemistry (phiPSII) in droughted compared with watered plants. However, phiPSII was larger at 0900 h than at 1400 h. The de-epoxidation state of the xanthophyll cycle increased with drought and was linearly related to NPQ. Intrinsic quantum yield of PSII (FV/FM) measured at dusk was also lower in severely stressed plants than in controls. Under maximum photosynthetic photon flux and high decarboxylation rates of organic acids, the D1 content in leaves of droughted plants showing maximal CAM activity was identical to the controls; increased drought decreased D1 content by more than 30%. Predawn samples had D1 contents similar to leaves sampled at peak irradiance, with no signs of recovery after 12 h of darkness. It is concluded that under mild water stress, early induction of CAM, together with an increased energy dissipation by the xanthophyll cycle, prevents net degradation of D1 protein; when water deficit is more severe, CAM and xanthophyll cycle capacities for energy dissipation decline, and net degradation of D1 proceeds.     

Effects of Exogenous Putrescine on Chlorophyll Fluorescence Imaging and Heat Dissipation Capacity in Cucumber (Cucumis sativus L.) Under Salt Stress

The objective of this study was to identify the effects of exogenous putrescine on photosynthetic performance and heat dissipation capacity in cucumber seedlings under salt stress. The stress of 75 mM NaCl for 7 days caused a significant decrease in net photosynthetic rate (P _N ). The experiment employed a chlorophyll fluorescence imaging technique and demonstrated that the maximal quantum yield of photosystem II photochemistry (Fv/Fm) and the actual photochemical efficiency of photosystem II (ΦPSII) were reduced by salt stress. Moreover, salt stress markedly reduced the photochemical quenching coefficient (qP) and non-photochemical quenching coefficient (qN), and significantly increased non-regulated heat dissipation (ΦNO). However, stressed plants supplied with exogenous putrescine exhibited higher P _N and ΦPSII, which indicated that putrescine can alleviate the detrimental effects on photosynthesis induced by salt stress. Putrescine sprayed on stressed plants significantly enhanced the regulated energy dissipation (ΦNPQ) and decreased ΦNO. Application of exogenous putrescine also changed the levels of xanthophyll cycle components and further enhanced the de-epoxidation state of xanthophyll cycle pigments under salt stress. Under control conditions, putrescine exerted little influence on the photosynthetic parameters in cucumber leaves. In conclusion, the application of exogenous putrescine may improve the heat dissipation capacity by promoting the xanthophyll cycle to reduce the damage caused by excess excitation energy, thus enhancing the salt tolerance of cucumber seedlings.     

Photosynthesis is improved by exogenous glycinebetaine in salt-stressed maize plants

The effects of exogenous application of glycinebetaine (GB) (10 m M ) on growth, leaf water content, water use efficiency, photosynthetic gas exchange, and photosystem II photochemistry were investigated in maize plants subjected to salt stress (50 and 100 m M NaCl). Salt stress resulted in the decrease in growth and leaf relative water content as well as net photosynthesis and the apparent quantum yield of photosynthesis. Stomatal conductance, evaporation rate, and water use efficiency were decreased in salt-stressed plants. Salt stress also caused a decrease in the actual efficiency of PSII ( Φ _PSII), the efficiency of excitation energy capture by open PSII reaction centers ( F _v'/ F _m'), and the coefficients of photochemical quenching ( q _P) but caused an increase in non-photochemical quenching (NPQ). Salt stress showed no effects on the maximal efficiency of PSII photochemistry ( F _v/ F _m). On the other hand, in salt-stressed plants, GB application improved growth, leaf water content, net photosynthesis, and the apparent quantum yield of photosynthesis. GB application also increased stomatal conductance, leaf evaporation rate, and water use efficiency. In addition, GB application increased Φ _PSII, F _v'/ F _m', and q _P but decreased NPQ. However, GB application showed no effects on F _v/ F _m. These results suggest that photosynthesis was improved by GB application in salt-stressed plants and such an improvement was associated with an improvement in stomatal conductance and the actual PSII efficiency.     

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。     

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.     

Leaf rolling and photosystem II efficiency in <Emphasis Type="Italic">Ctenanthe setosa</Emphasis> exposed to drought stress

Photochemical efficiency of PSII of Ctenanthe setosa was investigated to understand the photosynthetic adaptation mechanism under drought stress causing leaf rolling. Stomatal conductance (g _s), the levels of photosynthetic pigments and chlorophyll (Chl) fluorescence parameters were determined in leaves that had four different visual leaf rolling scores from 1 to 4, opened after re-watering and mechanically opened at score 4. g _s value gradually decreased in adaxial and abaxial surfaces in relation to scores of leaf rolling. Pigment contents decreased until score 3 but approached score 1 level at score 4. No significant variations in effective quantum yield of PSII (Φ_PSII), and photochemical quenching (q_p) were found until score 3, while they significantly decreased at score 4. Non-photochemical quenching (NPQ) increased at score 2 but then decreased. After re-watering, the Chl fluorescence and other physiological parameters reached to approximately score 1 value, again. As for mechanically opened leaves, g _s decreased during drought period. The decrease in adaxial surface was higher than that of the rolled leaves. NPQ was higher than that of the rolled leaves. Φ_PSII and q_p significantly declined and the decreases were more than those of the rolled leaves. In conclusion, the results indicate that leaf rolling protects PSII functionality from damage induced by drought stress.     

不同形态钙对高温逆境下番茄叶片光合作用的调控作

以‘辽园多丽’番茄幼苗为试材,在高温胁迫下喷施氯化钙(CaCl₂)、纳米钙(Nano-Ca)和糖醇钙(Manntiol-Ca),研究不同形态钙处理对高温逆境下番茄叶片光合作用的调控作用.结果表明: 不同形态的钙处理均可抑制高温逆境下叶片中叶绿素a和类胡萝卜素含量的下降;显著提高净光合速率(P_n),不同程度提高蒸腾速率(T_r)和气孔导度(g_s);降低PSII非调节性能量耗散的量子产量\[Y(NO)\]和由于受体侧限制引起的PSI非光化学能量耗散的量子产量\[Y(NA)\],提高PSII调节性能量耗散的量子产量\[Y(NPQ)\]和由于供体侧限制引起的PSI非光化学能量耗散的量子产量\[Y(ND)\];提高叶片中钙含量;Manntiol-Ca和Nano-Ca处理的总体效果优于CaCl₂处理.与CaCl₂相比,Manntiol-Ca和Nano-Ca是提高高温逆境下番茄叶片光合作用更有效的钙制剂.     

Photosynthetic properties of an Arabidopsis thaliana mutant possessing a defective PsbS gene

We describe the properties of npq4-9, a new mutant of Arabidopsis thaliana (L.) Heynh. with reduced nonphotochemical quenching (NPQ) capacity that possesses a single amino acid substitution in the PsbS gene encoding PSII-S, a ubiquitous pigment-binding protein associated with photosystem II (PSII) of higher plants. Growth, photosynthetic pigment contents, and levels of the major PSII antenna proteins were not affected by npq4-9. Although the extent of de-epoxidatin of violaxanthin to antheraxanthin plus zeaxanthin for leaves displaying the mutant phenotype equaled or exceeded that observed for the wild type, the relative effectiveness of de-epoxidized xanthophylls in promoting NPQ was consistently lower for the mutant. Energy partitioning in PSII was analyzed in terms of the competition for singlet chlorophyll a among the processes of fluorescence, thermal dissipation, and photochemistry. The key processes of NPQ and photochemistry in open PSII centers are represented by the relative in vivo rate constants kN and kP0, respectively. The magnitude of kP0 in normal leaves declined only slightly with increasing kN, consistent with localization of NPQ primarily in the antenna complex. Conversely, a highly significant linear decline in kP0 with increasing kN was observed for the mutant, consistent with a role for the PSII reaction center in the NPQ mechanism. Although the PSII absorption cross-section for white light was not significantly different relative to that of the wild type, PSII quantum yield was significantly lower in the mutant. The resulting lower capacity for linear electron transport in the mutant primarily affected reduction of terminal acceptors other than CO2. Parallel measurements of fluorescence and in vivo absorbance at 820 nm indicated a consistently higher steady-state level of reduction of PSII acceptors and accumulation of P700+ for the mutant. This suggests that inter-photosystem electron transport in the mutant is restricted either by a higher transthylakoid delta pH or by diminished accessibility to reduced plastoquinone.     

Exogenous proline induces soluble sugar accumulation and alleviates drought stress effects on photosystem II functioning of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> leaves

The effects of exogenous applied proline (Pro), on photosystem II (PSII) photochemistry of drought stressed (DS) 4-week old Arabidopsis thaliana plants, was studied by using chlorophyll (chl) fluorescence imaging. The maximum quantum yield of PSII photochemistry (F _v /F _m) in DS plants decreased significantly to 77% of that of the control value, suggesting that DS plants could not maintain PSII function, possibly due to accelerated photoinhibition of PSII. Free Pro and total soluble sugars (SS) increased, in response to DS. Exogenous foliar application of Pro by spraying, led to a remarkable increase in the accumulation of Pro and surprisingly also of SS. Both of them served to scavenge reactive oxygen species (ROS), as it was evident by the decreased lipid peroxidation level measured as malondialdehyde (MDA). DS plants sprayed with Pro showed a tolerance to photoinhibition, this indicated by F _v/F _m being close to values typical of healthy leaves by maintaining more than 98% of PSII function. Also the higher quantum efficiency of PSII photochemistry (Φ _PSΙΙ ) and the decreased excitation pressure (1 ? q _p ) recorded for stressed leaves with Pro, lead us to conclude that Pro appears to be involved in the protection of chloroplast structures by quenching ROS. The enhanced dissipation of excess light energy of PSII, in part accounts for the observed increased resistance to DS in A. thaliana leaves with Pro. Our data pointed out that Pro signalling interacts with SS signaling pathway and provided a new insight in Pro metabolism.     

Modifications in photosystem II photochemistry in senescent leaves of maize plants

Analyses of CO2 exchange and chlorophyll fluorescence were carried out to assess photosynthetic performance during senescence of maize leaves. Senescent leaves displayed a significant decrease in CO2 assimilatory capacity accompanied by a decrease in stomatal conductance and an increase in intercellular CO2 concentration. The analyses of fluorescence quenching under steady-state photosynthesis showed that senescence resulted in an increase in non-photochemical quenching and a decrease in photo-chemical quenching. It also resulted in a decrease in the efficiency of excitation energy capture by open PSII reaction centres and the quantum yield of PSII electron transport, but had very little effect on the maximal efficiency of PSII photochemistry. The results determined from the fast fluorescence induction kinetics indicated an increase in the proportion of QB-non-reducing PSII reaction centres and a decrease in the rate of QA reduction in senescent leaves. Theoretical analyses of fluorescence parameters under steady-state photosynthesis suggest that the increase in the non-photochemical quenching was due to an increase in the rate constant to thermal dissipation of excitation energy by PSII and that the decrease in the quantum yield of PSII electron transport was associated with a decrease in the rate constant of PSII photochemistry. Based on these results, it is suggested that the decrease in the quantum yield of PSII electron transport in senescent leaves was down-regulated by an increase in the proportion of QB-non-reducing PSII reaction centres and in the non-photochemical quenching. The photosynthetic electron transport would thus match the decreased demand for ATP and NADPH in carbon assimilation which was inhibited significantly in senescent leaves.Key words: Chlorophyll fluorescence, gas exchange, maize (Zea mays L.), photochemical and non-photochemical quenching, photosystem II photochemistry.      

High resolution imaging of photosynthetic activities of tissues, cells and chloroplasts in leaves

Through imaging of chlorophyll fluorescence, it is possible to produce parameterized fluorescence images that estimate the operating quantum efficiency of photosystem II (PSII) photochemistry and which can be used to reveal heterogeneous patterns of photosynthetic performance within leaves. The operating quantum efficiency of PSII photochemistry is dependent upon the effective absorption cross-section of the light-harvesting system of PSII and the photochemical capacity of PSII. The effective absorption cross-section is decreased by the process of down-regulation, which is widely thought to operate within the pigment matrices of PSII and which results in non-photochemical quenching of chlorophyll fluorescence. The photochemical capacity is non-linearly related to the proportion of PSII centres in the 'open' state and results in photochemical quenching of chlorophyll fluorescence. Examples of heterogeneity of the operating quantum efficiency of PSII photochemistry during the induction of photosynthesis in maize leaves and in the chloroplast populations of stomatal guard cells of a leaf of Tradescantia albifora are presented, together with analyses of the factors determining this heterogeneity. A comparison of the operating quantum efficiency of PSII photochemistry within guard cells and adjacent mesophyll cells of Commelina communis is also made, before and after stomatal closure through a change in ambient humidity.     

Up-regulation of the mitochondrial alternative oxidase pathway enhances photosynthetic electron transport under drought conditions

The aim of this study was to explore the role of the mitochondrial alternative oxidase (AOX) in the protection of photosynthesis during drought in wheat leaves. The relative water contents of water-replete and drought-exposed wheat plants were 97.2+/-0.3 and 75+/-2, respectively. Drought increased the amount of leaf AOX protein and also enhanced the rate of AOX-dependent O(2) uptake by the respiratory electron transport chain. The amount of the reduced, active form of the AOX protein was specifically increased by drought. The AOX inhibitor salicylhydroxamic acid (1 mM; SHAM) inhibited 70% of AOX activity in vivo in both water-replete and drought-exposed plants. Plants treated with SHAM were then exposed to low (100), high (350), or excess light (800 mumol photons m(-2) s(-1)) for 90 min. SHAM did not modify chlorophyll a fluorescence quenching parameters in water-replete controls after any of these treatments. However, while the maximal quantum yield of photosystem II (PSII) electron transport (F(v)/F(m)) was not affected by SHAM, the immediate quantum yield of PSII electron transport (Phi(PSII)) and photochemical quenching (qP) were gradually reduced by increasing irradiance in SHAM-treated drought-exposed plants, the decrease being most pronounced at the highest irradiance. Non-photochemical quenching (NPQ) reached near maximum levels in plants subjected to drought at high irradiance. However, a combination of drought and low light caused an intermediate increase in NPQ, which attained higher values when AOX was inhibited. Taken together, these results show that up-regulation of the respiratory AOX pathway protects the photosynthetic electron transport chain from the harmful effects of excess light.     

Photosynthetic acclimation to drought stress in Agave salmiana Otto ex Salm-Dyck seedlings is largely dependent on thermal dissipation and enhanced electron flux to photosystem I

Agave salmiana Otto ex Salm-Dyck, a crassulacean acid metabolism plant that is adapted to water-limited environments, has great potential for bioenergy production. However, drought stress decreases the requirement for light energy, and if the amount of incident light exceeds energy consumption, the photosynthetic apparatus can be injured, thereby limiting plant growth. The objective of this study was to evaluate the effects of drought and re-watering on the photosynthetic efficiency of A. salmiana seedlings. The leaf relative water content and leaf water potential decreased to 39.6 % and ?1.1 MPa, respectively, over 115 days of water withholding and recovered after re-watering. Drought caused a direct effect on photosystem II (PSII) photochemistry in light-acclimated leaves, as indicated by a decrease in the photosynthetic electron transport rate. Additionally, down-regulation of photochemical activity occurred mainly through the inactivation of PSII reaction centres and an increased thermal dissipation capacity of the leaves. Prompt fluorescence kinetics also showed a larger pool of terminal electron acceptors in photosystem I (PSI) as well as an increase in some JIP-test parameters compared to controls, reflecting an enhanced efficiency and specific fluxes for electron transport from the plastoquinone pool to the PSI terminal acceptors. All the above parameters showed similar levels after re-watering. These results suggest that the thermal dissipation of excess energy and the increased energy conservation from photons absorbed by PSII to the reduction of PSI end acceptors may be an important acclimation mechanism to protect the photosynthetic apparatus from over-excitation in Agave plants.     

Changes of photosynthetic traits in beech saplings (Fagus sylvatica) under severe drought stress and during recovery

In the context of an increased risk of extreme drought events across Europe during the next decades, the capacity of trees to recover and survive drought periods awaits further attention. In summer 2005, 4-year-old beech (Fagus sylvatica L.) saplings were watered regularly or were kept for 4 weeks without irrigation in the field and then re-watered again. Changes of plant water status, leaf gas exchange and Chl a fluorescence parameters, as well as alterations in leaf pigment composition were followed. During the drought period, stomatal conductance (g(s)) and net photosynthesis (P(n)) decreased in parallel with increased water deficit. After 14 days without irrigation, stomata remained closed and P(n) was almost completely inhibited. Reversible downregulation of PSII photochemistry [the maximum quantum efficiency of PSII (F(v)/F(m))], enhanced thermal dissipation of excess excitation energy and an increased ratio of xanthophyll cycle pigments to chlorophylls (because of a loss of chlorophylls) contributed to an enhanced photo-protection in severely stressed plants. Leaf water potential was restored immediately after re-watering, while g(s), P(n) and F(v)/F(m) recovered only partially during the initial phase, even when high external CO(2) concentrations were applied during the measurements, indicating lasting non-stomatal limitations. Thereafter, P(n) recovered completely within 4 weeks, meanwhile g(s) remained permanently lower in stressed than in control plants, leading to an increased 'intrinsic water use efficiency' (P(n)/g(s)). In conclusion, although severe drought stress adversely affected photosynthetic performance of F. sylvatica (a rather drought-sensitive species), P(n) was completely restored after re-watering, presumably because of physiological and morphological adjustments (e.g. stomatal occlusions).     

Response of the photosynthetic apparatus of cotton (Gossypium hirsutum) to the onset of drought stress under field conditions studied by gas-exchange analysis and chlorophyll fluorescence imaging.

The functioning of the photosynthetic apparatus of cotton (Gossypium hirsutum) grown during the onset of water limitation was studied by gas-exchange and chlorophyll fluorescence to better understand the adaptation mechanisms of the photosynthetic apparatus to drought conditions. For this, cotton was grown in the field in Central Asia under well-irrigated and moderately drought-stressed conditions. The light and CO(2) responses of photosynthesis (A(G)), stomatal conductance (g(s)) and various chlorophyll fluorescence parameters were determined simultaneously. Furthermore, chlorophyll fluorescence images were taken from leaves to study the spatial pattern of photosystem II (PSII) efficiency and non-photochemical quenching parameters. Under low and moderate light intensity, the onset of drought stress caused an increase in the operating quantum efficiency of PSII photochemistry (varphi(PSII)) which indicated increased photorespiration since photosynthesis was hardly affected by water limitation. The increase in varphi(PSII) was caused by an increase of the efficiency of open PSII reaction centers (F(v)'/F(m)') and by a decrease of the basal non-photochemical quenching (varphi(NO)). Using a chlorophyll fluorescence imaging system a low spatial heterogeneity of varphi(PSII) was revealed under both irrigation treatments. The increased rate of photorespiration in plants during the onset of drought stress can be seen as an acclimation process to avoid an over-excitation of PSII under more severe drought conditions.