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.     

Photosynthesis and PSII functionality of drought-resistant and drought-sensitive weeping lovegrass plants

World areas subject to drought are expected to increase under conditions of climate change. The purpose of this study is to clarify the response of grass species that can grow and produce under water stress. Therefore leaf photosynthesis, chlorophyll fluorescence and pigment content response to water stress were studied in two varieties of the C₄ grass Eragrostis curvula. Two-year-old plants of cv Ermelo and Consol were grown in plastic pots. Drought stress was imposed by withholding irrigation for 15 days and then rewatering for 5 days. During drought relative water content (RWC) decreased 65% in cv Ermelo, while lower reductions of RWC were observed in cv Consol. During the experiment in cv Ermelo increasing drought stress severity caused large decreases in photosynthetic rates, maximal PSII photochemical efficiency (F_V/F_M) and leaf pigment content. Cv Consol showed small variations in these parameters. Compared to cv Consol, after 15 days of drought, effective PSII quantum yield (Φ_II) was significantly lower in cv Ermelo. Reductions of Φ_II were related to significant reductions of open PSII energy capture efficiency (F′_V/F′_M). Photosynthetic response to increasing PPFD levels and to internal CO₂ concentration (C_i) were reduced by drought in cv Ermelo. Compared to well-watered control plants and to cv Consol, drought stressed plants of cv Ermelo showed also reductions of the initial slope of photosynthetic response to C_i and in the photosynthetic rate measured at saturating C_i. Moreover stomatal conductance (g) of both cvs decreased during drought. However, g was lower in drought stressed plants of cv Consol than in cv Ermelo. Water stress caused large reductions in leaf chlorophyll and carotenoid content in cv Ermelo, and small reductions in cv Consol. In drought-sensitive cv Ermelo water stress reduced the capabilities to down regulate PSII functionality through thermal energy dissipation. Results suggest that drought resistance of cv Consol, can be attributed to a higher water use efficiency.     

Changes in the photosynthetic efficiency of winter wheat in response to abiotic stress

The assessment of heat and drought tolerance is of primary importance in breeding programmes designed to improve heat and drought tolerance in cereals. Three winter wheat varieties grown in controlled growth chambers were exposed to heat (H) and drought (D) stress singly and in combination (H+D). The combined effects of H and D stress were much more severe than those of individual treatments for both physiological and yield parameters during grain filling. The chlorophyll content, effective quantum yield of PSII, net assimilation rate, transpiration, stomatal conductance and intercellular CO₂ concentration were greatly reduced by H, D and their interaction. Grain yield decreased to a greater extent (48.3%) in Plainsman V, averaged over the stress treatments, than in Mv Magma (67.8%) and Fatima 2 (53.7%). The least decline was found in grain number, except in Plainsman V. Mv Magma tolerated heat stress better than Fatima 2. In terms of photosynthetic activity, Plainsman V showed better drought tolerance than Mv Magma. The results showed that changes in physiological properties during stress treatment are not always associated with changes in yield parameters, so a combination of methods may be needed to give a more precise picture of the stress tolerance of wheat varieties.     

Alleviation of photoinhibition in drought-stressed wheat (Triticum aestivum) by foliar-applied glycinebetaine

Effects of foliar application of 100 mmol/L glycinebetaine (GB) on PS II photochemistry in wheat (Triticum aestivum) flag leaves under drought stress combined with high irradiance were investigated. The results show that GB-treated plants maintained a higher net photosynthetic rate during drought stress than non-GB treated plants. Exogenous GB can preserve the photochemical activity of PSII, for GB-treated plants maintain higher maximal photochemistry efficiency of PSII (F(v)/F(m)) and recover more rapidly from photoinhibition. In addition, GB-treated plants can maintain higher anti-oxidative enzyme activities and suffer less oxidative stress. Our data suggest that GB may protect the PSII complex from damage through accelerating D1 protein turnover and maintaining anti-oxidative enzyme activities at higher level to alleviate photodamage. Diethyldithiocarbamate as well as streptomycin treatment can impair the protective effect of GB on PSII. In summary, GB can enhance the photoinhibition tolerance of PSII.     

Comparative effect of water, heat and light stresses on photosynthetic reactions in Sorghum bicolor(L.) Moench

Five varieties of Sorghum bicolor (L.) Moench., differing in their drought tolerance under field conditions have been used to study the effect of individual components of drought stress, namely high light intensity stress, heat stress and water stress, on their photosynthetic performance. Chlorophyll content, chlorophyll fluorescence, ribulose-1,5-bisphosphate carboxylase (Rubisco, EC 4.1.1.39) content, phosphoenolpyruvate carboxylase (PEPcase, EC 4.1.1.31) activity and photo-synthetic oxygen evolution were used as key parameters to assess photosynthetic performance. The results indicated that photochemical efficiency of photosystem II (PSII) was severely reduced by all three stress components, whereas PEPcase activity was more specifically reduced by water stress. Degradation of Rubisco and chlorophyll loss occurred under high light and water stress conditions. Of the four drought-tolerant varieties, E 36-1 showed higher PEPcase activity, Rubisco content and photochemical efficiency of PSII, and was able to sustain a higher maximal rate of photosynthetic oxygen evolution under each stress condition as compared to the other varieties. A high stability to stress-induced damage, or acclimation of photosynthesis to the individual components of drought stress may contribute to the high yields of E 36-1 under drought conditions. In the E 36-1 variety markedly higher levels of the chloroplastic chaperonin 60 (cpn 60) were observed under all stress conditions than in the susceptible variety CSV 5.Key words: Chlorophyll fluorescence, drought stress, oxygen evolution, phosphoenopyruvate carboxylase, Sorghum.      

Photosystem II energy use,non-photochemical quenching and the xanthophyll cycle in Sorghumbicolor grown under drought and free-air CO2 enrichment(FACE) conditions

The present study was carried out to test the hypothesis thatelevated atmospheric CO₂ (Ca) will alleviate over‐excitationof the C₄ photosynthetic apparatus and decrease non‐photochemicalquenching (NPQ) during periods of limited water availability. Chlorophyll a fluorescencewas monitored in Sorghum bicolor plants grown under a free‐aircarbon‐dioxide enrichment (FACE) by water‐stress (Dry) experiment.Under Dry conditions elevated Ca increased the quantum yield ofphotosystem II (φPSII) throughout the day throughincreases in both photochemical quenching coefficient (q_p)and the efficiency with which absorbed quanta are transferred toopen PSII reaction centres (F_v′/F_m′).However, in the well‐watered plants (Wets) FACE enhanced φPSIIonly at midday and was entirely attributed to changes in F_v′/F_m′_. Underfield conditions, decreases in φPSII under Dry treatmentsand ambient Ca corresponded to increases in NPQ but the de‐epoxidation stateof the xanthophyll pool (DPS) showed no effects. Water‐stress didnot lead to long‐term damage to the photosynthetic apparatus asindicated by φPSII and carbon assimilation measuredafter removal of stress conditions. We conclude that elevated Caenhances photochemical light energy usage in C₄ photosynthesisduring drought and/or midday conditions. Additionally,NPQ protects against photo‐inhibition and photodamage. However,NPQ and the xanthophyll cycle were affected differently by elevatedCa and water‐stress.     

Effects of salicylic acid on protein kinase activity and chloroplast D1 protein degradation in wheat leaves subjected to heat and high light stress

In the north of China, wheat plants are often stressed by heat and high light during grain-filling stage, which leads to injury in photosynthetic apparatus and decline in photosynthetic rate. In order to develop a method to protect photosynthetic apparatus in wheat leaves subjected to heat and high light stress, the effects of SA (salicylic acid) and FSBA (5′-p-fluorosulfonylbenzoyl adenosine) on PK (protein kinase) activity, D1 protein degradation and the performance of PSII were investigated in present work. Our results showed that PK activity enhanced under heat and high light stress and declined when stress was removed. FSBA pretreatment resulted in marked decreases in PK activity and D1 protein level, suggesting a correlationship between degradation of D1 protein and phosphorylation. After 2 h of stress, D1 protein level in water-pretreated leaves decreased to 79% of control and then recovered to 81% after 3 h of recovery. This clearly indicated that the damage of D1 protein induced by heat and high light stress was reversible. Compared to the control, SA pretreatment could not only increase PK activity, retard the degradation of D1 protein during heat and high light stress, but also accelerate the recovery of D1 protein level when the stress was removed. Correspondingly, F_v/F_m (maximum photochemical efficiency of PSII), Φ_PSII (actual photochemical efficiency of PSII), ETR (electron transfer rate) and Pn (net photosynthetic rate) in SA-treated leaves were higher than that in leaves of control under both stress and non-stress conditions. Taken together, our results revealed that SA pretreatment could significantly alleviate damages of heat and high light stress on D1 protein and PSII of wheat leaves, and accelerate restoration of photosynthetic function.     

Stress-induced degradation of the photosynthetic apparatus is accompanied by changes in thylakoid protein turnover and phosphorylation

Development of chlorosis and loss of PSII were compared in young spinach plants suffering under a combined magnesium and sulphur deficiency. Loss of chlorophyll could be detected already after the first week of deficiency and preceded any permanent functional inhibition of PSII as detected by changes in the chlorophyll fluorescence parameter F_v/F_m. A substantial decrease in F_v/F_m was observed only after the second week of deficiency. After 4 weeks, the plants had lost about 70% of their original chlorophyll content, but fluorescence data indicated that 80% of the existing PSII centers were still capable of initiating photosynthetic electron transport. The degradation of the photosynthetic apparatus without loss of PSII activity was due to changes in protein turnover, especially of the PSII D1 reaction center protein. Already by day 7 of deficiency, a 1.4-fold increase in D1 protein synthesis was observed measured as incorporation of ¹⁴C-leucine. Immunological determination by western-blotting did not reveal a change in D1 protein content. Thus, D1 protein was also degraded more rapidly. The increased turnover was high enough to prevent any loss or inhibition of PSII. After 3 weeks, D1 protein synthesis on a chlorophyll basis was further increased by a factor of 2. However, this was not enough to prevent a net loss of D1 protein of about 70%. Immunological determination revealed that together with the D1 protein also other polypeptides of PSII became degraded. This process prevented a large accumulation of photo-inactivated PSII centers. However, it initiated the breakdown of the other thylakoid proteins, especially of LHCII, resulting in the observed chlorosis. Together with the change in protein turnover and stability, a characteristic change in thylakoid protein phosphorylation was observed. In the deficient plants steady state phosphorylation of both LHCII and PSII proteins was increased in the dark. In the light phosphorylation of PSII proteins was stimulated and after 3 weeks of deficiency was even higher in the deficient leaves than in the control plants. In contrast, the phosphorylation level of LHCII decreased in the light and could hardly be detected after 3 weeks of deficiency. Phosphorylation of the reaction center polypeptides presumably increased their stability against proteolytic attack, whereas phosphorylated LHCII seems to be the substrate for proteolysis.     

The increase in unsaturation of fatty acids of phosphatidylglycerol in thylakoid membrane enhanced salt tolerance in tomato

Overexpression of chloroplastic glycerol-3-phosphate acyltransferase gene (LeGPAT) in tomato increased cis-unsaturated fatty acid content in phosphatidylglycerol (PG) of thylakoid membrane. By contrast, suppressing the expression of LeGPAT decreased the content of cis-unsaturated fatty acid in PG. Under salt stress, sense transgenic plants exhibited higher activities of chloroplastic antioxidant enzymes, lower content of reactive oxygen species (ROS) and less ion leakage compared with the wild type (WT) plants. The net photosynthetic rate (P _N) and the maximal photochemical efficiency (F_v/F_m) of photosystem II (PSII) decreased more slightly in sense lines but more markedly in the antisense ones, compared to WT. D1 protein, located in the reactive center of the PSII, is the primary target of photodamage and has the highest turnover rate in the chloroplast. Under salt stress, compared with WT, the content of D1 protein decreased slightly in sense lines and significantly in the antisense ones. In the presence of streptomycin (SM), the net degradation of the damaged D1 protein was faster in sense lines than in other plants. These results suggested that, under salt-stress conditions, increasing cis-unsaturated fatty acids in PG by overexpression of LeGPAT can alleviate PSII photoinhibition by accelerating the repair of D1 protein and improving the activity of antioxidant enzymes in chloroplasts.     

Response of photosynthesis to high light and drought for Arabidopsis thaliana grown under a UV-B enhanced light regime

Arabidopsis thaliana grown in a light regime that included ultraviolet-B (UV-B) radiation (6 kJ m⁻² d⁻¹) had similar light-saturated photosynthetic rates but up to 50% lower stomatal conductance rates, as compared to plants grown without UV-B radiation. Growth responses of Arabidopsis to UV-B radiation included lower leaf area (25%) and biomass (10%) and higher UV-B absorbing compounds (30%) and chlorophyll content (52%). Lower stomatal conductance rates for plants grown with UV-B radiation were, in part, due to lower stomatal density on the adaxial surface. Plants grown with UV-B radiation had more capacity to down regulate photochemical efficiency of photosystem II (PSII) as shown by up to 25% lower φ_PSII and 30% higher non-photochemical quenching of chlorophyll fluorescence under saturating light. These contributed to a smaller reduction in the maximum photochemical efficiency of PSII (F _v/F _m), greater dark-recovery of F _v/F _m, and higher light-saturated carbon assimilation and stomatal conductance and transpiration rates after a four-hour high light treatment for plants grown with UV-B radiation. Plants grown with UV-B were more tolerant to a 12 day drought treatment than plants grown without UV-B as indicated by two times higher photosynthetic rates and 12% higher relative water content. UV-B-grown plants also had three times higher proline content. Higher tolerance to drought stress for Arabidopsis plants grown under UV-B radiation may be attributed to both increased proline content and decreased stomatal conductance. Growth of Arabidopsis in a UV-B-enhanced light regime increased tolerance to high light exposure and drought stress.     

24-epibrassionlide improves photosynthetic response of Rhododendron delavayi to drought

Rhododendron delavayi is an alpine evergreen ornamental plant with strong tolerance to drought stress. Brassinosteroids are promising agents for alleviating the negative effects of drought on plants, but the mechanism by which BRs induce plant resistance to drought is not well understood. The present study investigated the effects of exogenous spray of 24-epibrassionlide (EBR) at different concentrations (0~1 mg l⁻¹) on the physiological response of R. delavayi to drought caused by no watering for 10 days. With the increase in EBR concentration, net photosynthetic rate, stomatal conductance, transportation rate, light saturated photosynthetic rate, light compensation point, light saturation point, excitation energy capture efficiency of reaction center, actual photochemical efficiency of photosystem II (PSII), photochemical quenching and electron transport rate significantly increased, but there were no significant effects on photosynthetic pigment content. These results suggested that the EBR-induced improvement in CO₂ assimilation under drought was mainly related to stomatal and non-stomatal factors, and partially attributed to the increased photochemical efficiency of PSII. In addition, the leaf water potential increased with the increase in EBR concentration, while the malondialdehyde, superoxide dismutase, catalase, proline and soluble protein decreased. The results suggested EBR application partially alleviated the negative effect of drought on R. delavayi by improving water relations and decreasing lipid peroxidation and reactive oxygen species production. We concluded that exogenous application of EBR improved photosynthesis and alleviated the negative effects of drought-induced membrane peroxidation and severe oxidative stress.     

Photosynthetic response of sweet sorghum to drought and re‐watering at different growth stages

Sweet sorghum (Sorghum bicolor) is a C4 drought resistant species with a huge potential for bioenergy. Accentuated reductions in water availability for crop production and altered rainfall distribution patterns, however, will have direct impact on its physiological attributes, metabolic functions and plant growth. The objective of this study was to evaluate the effects of drought and re‐watering on the photosynthetic efficiency of sweet sorghum. Durable or short transient drought stress periods were imposed at early and late growth stages and compared with well‐watered plants. In spite of very similar drought levels at early and late growth stages (Ψ_soil = ?1.6 and ?1.7 MPa), the decrements in maximum quantum yield (?_Po) and performance index (PI) were about twice at late than at early growth stages. All the PI components, that is, density of active reaction centers (RCs), excitation energy trapping and conversion of excitation energy into electron flow followed a similar decreasing pattern. Upon re‐watering and regardless the duration and growth stage of the drought period, all the photosynthetic functions, and particularly those of photosystem II (PSII), fully recovered. Such effective self‐regulating functional activity by PSII photochemistry likely contributes to both high drought resistance and photosynthetic recovery capacity of sweet sorghum. At vegetative growth stages, the down regulation of the photochemistry seems to be the main photoprotective/regulative mechanisms, while at late growth stages, the accumulation of compatible solutes likely has a more preponderant role. The observed sugar concentration increments likely contributed to prevent permanent photo‐oxidative destruction of the PSII RCs of mature droughted sweet sorghum plants.     

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.     

The effect of CaCl<Subscript>2</Subscript> on calcium content,photosynthesis, and chlorophyll fluorescence of tung tree seedlings under drought conditions

The study investigated the effects of different CaCl₂ concentrations (2, 5, and 10 mM) on photosynthetic enzymatic activities, photosynthesis, and chlorophyll fluorescence of tung tree seedlings under drought conditions. Plants were sprayed with either CaCl₂ or distilled water until run-off. Irrigation was then withheld to induce drought stress. The strength of drought stress was evaluated by relative leaf water content and soil water content, which was 27.3 and 9.5% on day 0 and day 12, respectively. Drought stress decreased activities of ribulose-1,5-bisphosphate carboxylase/oxygenase and phosphoenolpyruvate carboxylase, chlorophyll (a+b) content, net photosynthetic rate, stomatal conductance, transpiration rate, electron transport rate, the maximal quantum yield of PSII photochemistry, and effective quantum yield of PSII in tung tree seedlings. The CaCl₂ pretreatments alleviated the negative effect of drought stress to some degree on all the parameters mentioned above.     

Water stress effects on photosynthesis in different mulberry cultivars

The effect of water stress on photosynthesis was determined in five mulberry cultivars (Morus alba L. cv. K-2, MR-2, BC2-59, S-13 and TR-10). Drought was imposed by withholding water and the plants were maintained at different water potentials ranging from 0.5 -MPa to 2.0 -MPa. Photosynthetic rates, activities of ribulose-1,5-bisphosphate carboxylase and sucrose phosphate synthase, photosystem II activity and chlorophyll content were used as key parameters to assess photosynthetic performance. There was a marked variation in the photosynthetic rates and ribulose-1,5-bisphosphate carboxylase activity among the five mulberry cultivars subjected to water stress. Photosystem II (PSII) and sucrose phosphate synthase activities were also severely reduced as measured by drought conditions. Of the five mulberry cultivars, S-13 and BC2-59 showed higher photosynthetic rates, ribulose-1,5-bisphosphate carboxylase activity, high sucrose phosphate synthase activity and photochemical efficiency of PSII compared to the other varieties.     

Effects of four types of sodium salt stress on plant growth and photosynthetic apparatus in sorghum leaves

Sorghum variety Longza 17 was used as the experimental organism in a study of the effects of different types of sodium salt (two neutral salts, NaCl and Na₂SO₄; and two alkaline salts, NaHCO₃ and Na₂CO₃), at an equivalent Na⁺ concentration (100?mmol·L^?1) on leaf growth parameters and PSII and PSI function by using the Fast Chlorophyll Fluorescence Induction Dynamics technique and 820?nm light reflectance curves. The results showed that at Na⁺ concentration of 100?mmol·L^?1, different types of sodium salt stress significantly inhibited the growth of sorghum plants. Different types of sodium salt stress showed significant inhibition on the activities of PSII and PSI in sorghum leaves, the impact of different types of sodium salt on the activities of PSII and PSI in sorghum leaves was consistent, listed from greatest to least impact as Na₂CO₃ > NaHCO₃ > Na₂SO₄ > NaCl. The effects of alkaline salt stress on the growth and photosynthetic properties of sorghum were greater than those under the neutral salt stress, therefore, in addition to considering the impact of Na⁺ concentration in the sorghum planting area, emphasis should also be given to the influence of the degree of alkalization, especially the higher alkalinity of Na₂CO₃.     

Photosynthetic electron transport and specific photoprotective responses in wheat leaves under drought stress

The photosynthetic responses of wheat (Triticum aestivum L.) leaves to different levels of drought stress were analyzed in potted plants cultivated in growth chamber under moderate light. Low-to-medium drought stress was induced by limiting irrigation, maintaining 20 % of soil water holding capacity for 14 days followed by 3 days without water supply to induce severe stress. Measurements of CO₂ exchange and photosystem II (PSII) yield (by chlorophyll fluorescence) were followed by simultaneous measurements of yield of PSI (by P700 absorbance changes) and that of PSII. Drought stress gradually decreased PSII electron transport, but the capacity for nonphotochemical quenching increased more slowly until there was a large decrease in leaf relative water content (where the photosynthetic rate had decreased by half or more). We identified a substantial part of PSII electron transport, which was not used by carbon assimilation or by photorespiration, which clearly indicates activities of alternative electron sinks. Decreasing the fraction of light absorbed by PSII and increasing the fraction absorbed by PSI with increasing drought stress (rather than assuming equal absorption by the two photosystems) support a proposed function of PSI cyclic electron flow to generate a proton-motive force to activate nonphotochemical dissipation of energy, and it is consistent with the observed accumulation of oxidized P700 which causes a decrease in PSI electron acceptors. Our results support the roles of alternative electron sinks (either from PSII or PSI) and cyclic electron flow in photoprotection of PSII and PSI in drought stress conditions. In future studies on plant stress, analyses of the partitioning of absorbed energy between photosystems are needed for interpreting flux through linear electron flow, PSI cyclic electron flow, along with alternative electron sinks.     

Sugar accumulation,photosynthesis and growth of two indica rice varieties in response to salt stress

Sugar, a final product of photosynthesis, is reported to be involved in the defense mechanisms of plants against abiotic stresses such as salinity, water deficiency, extreme temperature and mineral toxicity. Elements involved in photosynthesis, sugar content, water oxidation, net photosynthetic rate, activity of enzyme and gene expression have therefore been studied in Homjan (HJ), salt-tolerant, and Pathumthani 1 (PT1), salt-sensitive, varieties of rice. Fructose-1,6-biphosphatase (FBP) and fructokinase (FK) genes were rapidly expressed in HJ rice when exposed to salt stress for 1–6 h and to a greater degree than in PT1 rice. An increase in FBP enzyme activity was found in both roots and leaves of the salt-tolerant variety after exposure to salt stress. A high level of sugar and a delay in chlorophyll degradation were found in salt-tolerant rice. The total sugar content in leaf and root tissues of salt-tolerant rice was 2.47 and 2.85 times higher, respectively, than in the salt-sensitive variety. Meanwhile, less chlorophyll degradation was detected. Salt stress may promote sugar accumulation, thus preventing the degradation of chlorophyll. Water oxidation by the light reaction of photosynthesis in the salt-tolerant variety was greater than that in the salt-sensitive variety, indicated by a high maximum quantum yield of PSII (F _v/F _m) and quantum efficiency of PSII (Φ_PSII) with low nonphotochemical quenching (NPQ), leading to a high net photosynthetic rate. In addition, the overall growth performances in the salt-tolerant variety were higher than those in the salt-sensitive variety. The FBP gene expression and enzyme activity, sugar accumulation, pigment stabilization, water oxidation and net photosynthetic rate parameters in HJ rice should be further investigated as multivariate salt-tolerant indices for the classification of salt tolerance in rice breeding programs.     

Drought effect on photosynthetic activity,osmolyte accumulation and membrane integrity of two <Emphasis Type="Italic">Cicer arietinum</Emphasis> genotypes

Drought was induced in chickpea (Cicer arietinum L.) genotypes (ChK 3226 and ILC 3279) differing in yield capacity. Water stress (S1, RWC around 55–50%; S2, RWC ≤ 40%) drastically reduced stomatal conductance (g _s) and net photosynthetic rate (P _N) in both genotypes. ILC 3279 showed greater photosynthetic capacity (A _max) decreases. Maximum PSII photochemical efficiency (F_v/F_m), photochemical quenching (q_P), total chlorophylls (Chls) and carotenoids (Cars) content showed stability in both genotypes under stress, but in S2 ILC 3279 presented an increase in basal fluorescence (F₀) and a greater reduction in estimation of quantum yield of linear electron transport (Φ_e) than ChK 3226. Membrane damage evaluated by electrolyte leakage occurred earlier and was greater in ILC 3279. It also presented a decrease of total fatty acids (TFA) along drought, while in ChK 3226 greater amounts of TFA were observed in S1. In rehydration, P _N of S1 plants completely recovered (ILC 3279) or remained slightly below control (ChK 3226). As regards S2 plants, ILC 3279 showed stronger P _N and g _s reductions than ChK 3226, despite both genotypes totally recovered A _max and chlorophyll (Chl) a fluorescence. ChK 3226 recovered more efficiently from membrane damage. Under control conditions, greater amounts of most of the studied soluble metabolites occurred in ChK 3226 plants. Malate and citrate decreased with water stress (S2) in both genotypes. Sucrose and pinitol (that had a higher concentration than sucrose in both genotypes) increased in ILC 3279 (S1 and S2), and decreased in ChK 3226 (S2). In ILC 3279 proline and asparagine followed similar patterns. Genotypes showed a similar shoot dry mass (DM) in control plants, but root DM was higher in ChK 3226. Drought reduced root and shoot DM in ChK 3226 already under S1, while in ILC 3279 root DM was unaffected by drought and shoot biomass decreased only in S2. Root/shoot ratio was always higher in ChK 3226 but tended to decrease under stress, while the opposite was observed in ILC 3279. No pods were obtained from control plants of both genotypes, or droughted ILC 3279 plants. ChK 3226 produced pods under S1 (higher yield) and S2. Under stress conditions, ChK 3226 was less affected in photosynthetic activity and membrane integrity, showing a better tolerance to drought. This agrees with the better yield of this genotype under water stress. Distinct strategies seem to underlie the different physiological responses of the two genotypes to water deficit. In spite of its significant solutes accumulation, ILC 3279 was more affected in photosynthetic activity and membrane integrity during water stress than ChK 3226, which showed better yield under drought. A relation could not be established between solutes accumulation of ILC 3279 and yield.     

Effects of Drought Stress on Key Enzymes of Carbon Metabolism,Photosynthetic Characteristics and Agronomic Traits of Soybean at the Flowering Stage under Different Soil Substrates

Soybean is an important legume food crop, and its seeds are rich in nutrients, providing humans and animals with edible oil and protein feed. However, soybean is sensitive to water requirements, and drought is an important factor limiting soybean yield and quality. This study used Heinong 84 (drought resistant variety) and Hefeng 46 (intermediate variety) as tested varieties planted in chernozem, albic, and black soils. The effects of drought stress on the activities of key enzymes in carbon metabolism and photosynthetic characteristics of soybean were studied during the flowering stage, most sensitive to water. (1) The activities of SS-1, 6PGDH, and G6PDH enzymes in soybean leaves first increased and then decreased under drought stress. The enzyme activity was the highest under moderate drought stress and weakest in the blank group. (2) Drought stress increased Phi2, PhiNO, and Fm in soybean leaves and reached the highest value under severe drought; with the increase in drought stress, PhiNPQ and Fv/Fm of soybean leaves gradually decreased, reaching the lowest under severe drought. (3) With the increase in drought stress, F₀ and Fs of soybean leaves showed a single peak curve, and the maximum was at moderate drought. (4) Correlation analysis showed that F₀ was greatly affected by varieties and soil types; Fs, F0, and Fm soil varieties had a great influence, and chlorophyll fluorescence parameters were affected differently under drought stress with different drought degrees. (5) Drought stress changed the agronomic traits and yield of soybean. With the increase of drought degree, plant height, node number of main stem, effective pod number, 100-seed weight and total yield decreased continuously. (6) Drought stress affected the dry matter accumulation of soybean. With the increase of drought degree, the dry matter accumulation gradually decreased. Among them, the leaf was most seriously affected by drought, and SD decreased by about 55% compared with CK. Under the condition of black soil, the dry matter accumulation of soybean was least affected by drought.