Water Stress Induced Proline Accumulation in Contrasting Wheat Genotypes as Affected by Calcium and Abscisic Acid

Proline accumulation and mobilization in roots of 7-d-old seedlings of wheat genotypes varying in sensitivity towards water stress were compared. Water stress was induced by polyethylene glycol (PEG-6000; osmotic potential −1.5 MPa) in the presence of 0.1 mM abscisic acid (ABA), 1 mM calcium chloride, 0.5 mM verapamil (Ca²⁺ channel blocker), 0.5 mM fluridone (inhibitor of ABA biosynthesis). While both the genotypes did not differ in total proline accumulation, rate of proline accumulation and utilization was higher in tolerant genotype C 306 as compared to susceptible genotype HD 2380. The treatment with ABA and CaCl₂ caused further increase in proline accumulation during stress and reduced its mobilization during recovery. The membrane stability and elongation rate of roots was observed to be higher at ABA and calcium treatment in both the genotypes under stress. As was evident from inhibitor studies, the tolerant genotype was more responsive to ABA and the susceptible one to calcium. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photosynthetic Characteristics in Two Wheat Genotypes as Affected by Nitrogen Nutrition

Chlorophyll (Chl) a and b content, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) content and activity, and electron transport rate were measured in flag leaves of wheat genotypes Uniculm and Kalyansona, grown at suboptimal and optimal supply of nitrogen. The Chl content, RuBPCO activity, and electron transport rate were decreased due to suboptimal nitrogen supply only in Kalyansona. There was no change in the ratio of RuBPCO and photosystem 2 (PS2) activity at various stages which suggests that there was no alteration in distribution of N due to additional N supply. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Photosynthetic Traits in Wheat Grown under Decreased and Increased CO2 Concentration, and after Transfer to Natural CO2 concentration

Wheat plants were grown from sowing to day 18 in 26-dm³ chambers at three different CO₂ concentrations: 150 (-CO₂), 350 (C, control), 800 (+CO₂) mol mol^-1. Afterwards, plants of the three variants were grown at the same natural CO₂ concentration. Plant characteristics were measured just before the transfer (0 days after CO₂ treatment, DAT), and at 5 – 8 DAT on the 1^st leaf, and at 12 – 22 DAT on the 4^th leaf. Decreased or increased CO₂ concentrations caused acclimations which persisted after transplantation to natural CO₂ concentration. At 5 – 8 DAT, stomatal density, stomatal conductance (g_s), CO₂ saturated net photosynthetic rate (P_Nsat0), radiation saturated net photosynthetic rate (P_Nsat1), and carboxylation efficiency () were higher in -CO₂ plants and lower in +CO₂ plants than in C plants. As compared with C plants, the photochemical efficiency () was lower in -CO₂ and higher in -CO₂ plants, however, chlorophyll (Chl) a, Chl b, Chl a–b and carotenoid contents were lower in both -CO₂ and +CO₂ plants. On the 4^th leaf, which emerged on plant after finishing CO₂ treatments, at 12 – 22 DAT, no differences in stomatal density and g, between treatments were observed. In -CO₂ plants, pigment content and P_Nsat0 were higher, was lower, and P_Nsat1 and were not different from C plants. In contrast, in +CO₂ plants, pigment content, P_Nsat1 and were lower, and P_Nsat0 and were unchanged. Leaf area, dry mass, and tiller development increased in +CO₂ plants and decreased in -CO₂ plants. In the interval between 8 and 22 DAT, lower net assimilation rate in +CO₂ than in -CO₂ plants was observed.     

Photosynthetic carbon fixation in the corollas of Petunia hybrida

Corollas of Petunia hybrida (cv. Hit Parade Rosa) flowers fixed ¹⁴CO₂ under both light and dark conditions. Rates of light fixation were much higher in mature pink corollas than in young, green corollas [57 and 9 nmol (ngchl)¹ min^-1], paralleling the development of chloroplasts in these tissues. Stomatal conductance in corollas was only 12% of that in green leaves, mainly due to the presence of few, and non-functioning stomata in the corolla. The activity and concentration of ribulose bisphosphate carboxylase (EC 4.1.1.39) in corolla extracts were only about 30% (per unit Chi) of those in extracts from green leaves. These results, together with previous results, might indicate a coordinated reduction in activity of systems participating in photosynthesis in corollas. The fixation products following a 6 s pulse with ¹⁴CO₂, were typical of C, plants in both corollas and green leaves, but a higher level of β-carboxylation products was found in the corollas. The activity of phosphoenol-pyruvate carboxylase (EC 4.1.1.31) (per unit protein) was similar in both tissues. Although the total carbon fixed by the corolla constituted only a small part of the metabolites required for flower development, certain photosynthetic metabolites might have a regulatory role in flower development.     

Diurnal Oscillation in the Intercellular CO2 Concentration of Spring Wheat Under the Semiarid Conditions

Yields of wheat in semiarid and arid zones are limited by drought, and water condition is very important at each stage of development. Studies carried out at Loess Plateau in the northwestern part of China indicated that yield of spring wheat (Triticum aestivum L.) cv. Dingxi 81-392 was reduced by 41% when subjected to water stress. The effects of two water regimens on net photosynthetic rate (P _N), stomatal conductance (g _s), and intercellular CO₂ concentration (C _i) were investigated at the jointing, booting, anthesis, and grain filling stages. Low soil moisture in comparison to adequate one had invariably reduced P _N during the diurnal variations at the four growth stages. P _N and g _s in both soil moisture regimes was maximally reduced at midday. C _i and the stomatal limitation fluctuated remarkably during photosynthesis midday depression processes, especially at the grain filling stage. Hence atmospheric drought at midday was one of the direct causes inducing stomata closure and the g _s depression, but it was beneficial for maintaining stable intrinsic water use efficiency. Fluctuation in C _i implicated that non-stomatal limitation also plays an important role during the period of photosynthesis midday depression. Consequently stomatal and/or non-stomatal limitation are the possible cause of the midday photosynthesis decline.     

Effect of Water Stress on Photosynthesis and Growth in Two Teak Phenotypes

Two teak (Tectona grandis L.f.) phenotypes differing in their leaf length/breadth ratios were subjected to water stress by withholding water supply for three weeks. Growth rates of whole plants, developing leaves (1^st and 2^nd from shoot apices), and 2^nd and 3^rd internodes were higher in broad leaved (BL) phenotype than in narrow leaved (NL) phenotype before and after imposing water stress treatment. However, the effect of water stress on these parameters was higher in the BL phenotype than in the NL one. Diurnal course of net photosynthetic rate (P _N) of 3^rd or 4^th leaves from shoot apices measured under well-watered conditions was higher for the NL than BL phenotype. P _N, stomatal conductance (g _s), and transpiration rate (E) in both phenotypes were negatively affected by water stress and their decline under water stress was significantly higher in the BL than NL plants. This revised version was published online in June 2006 with corrections to the Cover Date.     

Hormonal Regulation of Photosynthetic Enzymes in Cotton under Water Stress

Activities of ribulose bisphosphate carboxylase/oxygenase (RuBPCO), phosphoenolpyruvate carboxylase (PEPC), and carbonic anhydrase (CA) were determined in leaves of cotton (Gossypium hirsutum L. cv. H-777) subjected to 8-d waterlogging (WL) at the vegetative stage, or to drought (D) at the reproductive stage, or to interaction of both stresses. The soil moisture of control plants was kept at field capacity. One day prior to stress various growth hormones (5 μM) were sprayed up to runoff. WL reduced RuBPCO and CA activities, while PEPC activity increased. Upon D, RuBPCO and PEPC activities were reduced while CA activity was increased. Imposition of both stresses increased activities of all three enzymes. Effect of stresses on enzyme activity was alleviated by benzylaminopurine (BAP), but indol-3-yl-acetic acid was more promoting under interactive stress. No CA activity with BAP was observed during interactive stress. This revised version was published online in August 2006 with corrections to the Cover Date.     

Modelling Net Photosynthetic Rate of Winter Wheat in Elevated Air CO2 Concentrations

Winter wheat plants were grown in open top chambers either at 365 µmol mol^–1 (AC) or at 700 µmol mol^–1 (EC) air CO₂ concentrations. The photosynthetic response of flag leaves at the beginning of flowering and on four vertical leaf levels at the beginning of grain filling were measured. Net photosynthetic rates (P _N) were higher at both developmental phases in plants grown at EC coupled with larger leaf area and photosynthetic pigment contents. The widely accepted Farquhar net photosynthesis model was parameterised and tested using several observed data. After parameterisation the test results corresponded satisfactorily with observed values under several environmental conditions.     

Interaction of Cytokinins and Abscisic Acid During Regulation of Stomatal Opening in Bean Leaves

Effects of benzyladenine (BA) and abscisic acid (ABA) applied separately or simultaneously on parameters of gas exchange of Phaseolus vulgaris L. leaves were studied. In the first two experimental sets) 100 M ABA and 10 M BA were applied to plants sufficiently supplied with water. Spraying of leaves with ABA decreased stomatal conductance (g _s) and in consequence transpiration rate (E) and net photosynthetic rate (P _N) already 1 h after application, but 24 h after application the effect almost disappeared. 10 M BA slightly decreased gas exchange parameters, but in simultaneous application with ABA reversed the effect of ABA. Immersion of roots into the same solutions markedly decreased gas exchange parameters and 24 h after ABA application the stomata were completely closed. The effect of ABA was ameliorated by simultaneous BA application, particularly after 1-h treatment. In the third experimental set, plants were pre-treated by immersing roots into water, 1 M BA, or 100 M ABA for 24 h and then the halves of split root system were dipped into different combinations of 1 M BA, 100 M ABA, and water. In plants pre-treated with ABA all gas exchange parameters were small and they did not differ in plants treated with H₂O+H₂O, H₂O+BA, or BA+BA. In plants pre-treated with BA or H₂O, markedly lower values of P _N were found when both halves of roots were immersed in ABA. Further, the effects of pre-treatment of plants with water, 1 M BA, 100 M ABA, or ABA+BA on the development of water stress induced by cessation of watering and on the recovery after rehydration were followed. ABA markedly decreased gas exchange parameters at the beginning of the experiment, but in its later phase the effect was compensated by delay in development of water stress. BA also delayed development of water stress and increased P _N in water-stressed leaves. BA reversed the effect of ABA at mild water stress. Positive effects of BA and ABA pre-treatments were observed also after rehydration.     

Effects of Pre-Treatments with Abscisic Acid and/or Benzyladenine on Gas Exchange of French Bean, Sugar Beet, and Maize Leaves During Water Stress and After Rehydration

Net photosynthetic rate (P_N), transpiration rate (E), and stomatal conductance (g_s) during water stress and after rehydration were measured in Phaseolus vulgaris, Beta vulgaris, and Zea mays. Immediately before imposition of water stress by cessation of watering, plants were irrigated with water (control), 100 M abscisic acid (ABA), and/or 10 M N⁶-benzyladenine (BA). In all three species, application of ABA decreased g_s, E, and P_N already 1 h after application. However, during water stress g_s, E, and P_N in plants pre-treated with ABA remained higher than in plants pre-treated with water. Positive effects of ABA application were observed also after rehydration. In contrast, the effects of pre-treatment with BA were species-specific. While in bean plants BA application ameliorated negative effect of water stress, only very slight effects were observed in maize, and in sugar beet BA even aggravated the effects of water stress.     

Assessment of Genotypic Variation in Salt Tolerance of early CIMMYT Hexaploid Wheat Germplasm Using Photosynthetic Capacity and Water Relations as Selection Criteria

Twenty-five genotypes of early CIMMYT hexaploid wheat were screened for salt tolerance in a glasshouse experiment using photosynthetic capacity and water relation parameters as selection criteria. Under salt stress (150 mM NaCl) the genotypes Frontana, Norin-10, Mayo-54, Noreste-66, and Yaktana-54 excelled all other lines in shoot dry mass, and Na(20)TPP, Penjamo-62, Inia-66, Frontana, Siete Cerros, and Jaral-66 in grain yield per plant in both absolute and relative (percent of control) terms. Although net photosynthetic rate (P _N) declined in all genotypes due to salt stress, it was not helpful in discriminating among genotypes according to salt tolerance. Similarly, no positive relationships of salt tolerance of the genotypes with stomatal conductance, leaf water potential, or turgor pressure were found. Every genotype used its own specific mechanism to tolerate salt stress. However, a large amount of variation in salt tolerance observed in 25 early CIMMYT wheat genotypes can be of considerable practical value for improving salt tolerance in the existing commercial hexaploid wheats. This revised version was published online in August 2006 with corrections to the Cover Date.     

Photosynthetic Enzymes of the C4 Grass Setaria sphacelata Under Water Stress: A Comparison Between Rapidly and Slowly Imposed Water Deficit

Two stress imposing systems were used: a rapid stress developed by allowing excised leaves to loose water by transpiration, and a slow stress developed by withholding watering of potted plants. Carboxylating enzymes reacted differently on both types of stress. Rapid stress increased ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activation, but both activities (initial and total) showed little variation with stress. Under slow stress the activation did not change, although both activities decreased much under stress. Phosphoenolpyruvate carboxylase (PEPC) showed a deep decrease of activity under rapid stress, nevertheless, a certain recovery was found under extreme stress. On the other hand, under slow stress the activity of PEPC showed a linear increase with decreasing relative water content. The ratio between physiological and maximal activity increased slightly under both types of stress. The activity of malic enzyme did not change under rapid stress, and decreased linearly under slow stress.     

Developmental and Photosynthetic Characteristics of a Photoautotrophic Chrysanthemum Culture

Chrysanthemum plantlets were cultivated in vitro on media with 2.0, 0.3, or 0 % sucrose, or photoautotrophically without an organic carbon source but with supplementation of the culture vessel atmosphere with 2 % CO₂. The photoautotrophically cultivated plantlets showed a better growth and multiplication, higher contents of chlorophyll (Chl) and carotenoids, higher Chl a/b ratio, net photosynthetic rate and ribulose-1,5-bisphosphate carboxylase/oxygenase and phosphoenolpyruvate carboxylase activities than plantlets grown on the medium with sucrose. This revised version was published online in June 2006 with corrections to the Cover Date.     

Plant Responses to Drought,Acclimation, and Stress Tolerance

At the whole plant level, the effect of stress is usually perceived as a decrease in photosynthesis and growth. That is why this review is focused mainly on the effect of drought on photosynthesis, its injury, and mechanisms of adaptation. The analysed literature shows that plants have evolved a number of adaptive mechanisms that allow the photochemical and biochemical systems to cope with negative changes in environment, including increased water deficit. In addition, the acquisition of tolerance to drought includes both phenotypic and genotypic changes. The approaches were made to identify those metabolic steps that are most sensitive to drought. Some studies also examined the mechanisms controlling gene expression and putative regulatory pathways.     

Chilling Induced Oxidative Stress in Germinating Wheat Grains as Affected by Water Stress and Calcium

Wheat (Triticum aestivum L.) plants were subjected to mild water stress during grain filling at milk (early, medium, and late) and dough (early, soft, hard) stages. The grains harvested from stressed plants were subjected to low temperature stress of 10 °C for 24 h in presence or absence of 1 mM CaCl₂, and embryos were examined for oxidative injury. The embryos of grains water stressed at milk and soft dough stages showed lowest contents of H₂O₂ and malondialdehyde and highest membrane stability index, ascorbic acid content, and activities of catalase, ascorbate peroxidase, and superoxide dismutase as compared to control embryos or water-stressed at other stages. Presence of Ca²⁺ in the medium reduced H₂O₂ and malondialdehyde content and increased ascorbic acid content, and catalase, ascorbate peroxidase and superoxide dismutase activities.     

Synergistic effects of cadmium and NaCl on the growth,photosynthesis and ion content in wheat plants

The addition of NaCl to cadmium had significant synergistic effect on the wheat root and shoot fresh mass, relative growth rate and net assimilation rate, while showed no significant effects on the dry mass production, leaf area, leaf area ratio, leaf mass ratio and specific leaf area. Additive depression of the rate of photosynthesis and the stomatal conductance was recorded, while no significant effect on the transpiration rate was observed. The Cd stress disturbed the mineral nutrition of the wheat plants either directly or indirectly, NaCl markedly reduce the uptake and internal concentration of K and Ca in the shoot. The combination of cadmium and NaCl showed no additive effects on the content of ions in the root as well as in the shoot of wheat plants.     

Changes of Photosynthetic Characteristics in Relation to Leaf Senescence in Two Maize Hybrids with Different Senescent Appearance

A field experiment was conducted to investigate the changes in chlorophyll (Chl) and nitrogen (N) contents, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and phosphoenolpyruvate carboxylase (PEPC) contents and PEPC activity, and the photon-saturated net photosynthetic rate (P _Nsat), and their relationships with leaf senescence in two maize hybrids with different senescent appearance. One stay-green (cv. P3845) and one earlier senescent (cv. Hokkou 55) hybrid were used in this study, and we found that Chl and N contents and the P _Nsat in individual leaves of P3845 were greater than those in corresponding leaves of Hokkou 55 at the successive growth stages. In addition, larger contents of RuBPCO and PEPC, and a greater activity of PEPC were observed in P3845. Due to the lower rates of decrease of Chl, RuBPCO, and PEPC amounts per unit of N, and the lower net C translocation rate per unit of N in the stay-green hybrid, leaf senescence was delayed in comparison to the earlier senescent hybrid.     

Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase Content and Activity in Wheat, Rye and Triticale

Photosynthetic parameters were measured in triticale and its parents wheat and rye. Soluble protein content in leaves, ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) content per fresh mass, total chlorophyll content, biomass yield, leaf area, leaf mass and specific leaf mass were higher but Rubisco content expressed as percentage of soluble protein, carboxylase activity, photosynthetic rate and stomatal conductance were significantly lower in rye than in wheat. Native-PAGE of Rubisco revealed that rye carboxylase was different from that of wheat. The difference was not related to either the small or large subunit of Rubisco but, may be, to the ionic and/or other properties of the Rubisco protein moiety. Triticale Rubisco was similar to wheat. For most of the studied physiological parameters, triticale showed much more similarity with wheat than with rye.     

Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought

A gradual reduction in leaf water potential (Ψ_leaf), net photosynthetic rate (P _N), stomatal conductance, and transpiration rate was observed in two drought tolerant (C 306 and K 8027) and two susceptible (RW 893 and 899) genotypes subjected to water stress. The extent of reduction was lower in K 8027 and C 306 and higher in RW 893 and RW 899. Rewatering the plants after 5 d of stress restored P _N and other gas exchange traits in all four cultivars. Water stress had no significant effect on variable to maximum fluorescence ratio (F_v/F_m) indicating that water stress had no effect on primary photochemistry of photosystem 2 (PS2). However, water stress reduced the efficiency of excitation energy transfer (F′_v/F′_m) and the quantum yield of electron transport (Φ_PS2). The reduction was more pronounced in susceptible cultivars. Water stress had no significant effect on photochemical quenching, however, the non-photochemical quenching increased by water stress.