Acclimation to long-term water deficit in the leaves of two sunflower hybrids: photosynthesis, electron transport and carbon metabolism

The influence of long-term water deficit on photosynthesis, electron transport and carbon metabolism of sunflower leaves has been examined. Water deficit was imposed from flower bud formation up to the stage of full flowering in the field on two sunflower hybrids with different drought tolerance. CO2 assimilation and stomatal conductance of the intact leaves, determined at atmospheric CO2 and full sunlight (1500-2000 mol quanta m^-2 s^-1), decreased with water deficit. Maximum quantum efficiency of PSII (Fy/Fm) and relative quantum yield of PSII (II) determined under similar experimental conditions, did not change significantly in severely stressed leaves. The strong inhibition of the plateau region of the light response curve, determined at high CO2 (5%) in water-deficient sunflower leaves, indicates that photosynthesis is also limited by non-stomatal factors. The decreased slope and the plateau of the CO2 response curves show that the capacity of carboxylation and RuBP regeneration decreased in severely stressed intact leaves. Rubisco specific activity decreased in severely stressed leaves, but Rubisco content increased under prolonged drought. The increase of Rubisco content was significantly higher in leaves of the drought-tolerant sunflower hybrid indicating that a higher Rubisco content could be one factor in conferring better acclimation and higher drought tolerance.     

Responses of leaf stomatal density to water status and its relationship with photosynthesis in a grass

Responses of plant leaf stomatal conductance and photosynthesis to water deficit have been extensively reported; however, little is known concerning the relationships of stomatal density with regard to water status and gas exchange. The responses of stomatal density to leaf water status were determined, and correlation with specific leaf area (SLA) in a photosynthetic study of a perennial grass, Leymus chinensis, subjected to different soil moisture contents. Moderate water deficits had positive effects on stomatal number, but more severe deficits led to a reduction, described in a quadratic parabolic curve. The stomatal size obviously decreased with water deficit, and stomatal density was positively correlated with stomatal conductance (g(s)), net CO(2) assimilation rate (A(n)), and water use efficiency (WUE). A significantly negative correlation of SLA with stomatal density was also observed, suggesting that the balance between leaf area and its matter may be associated with the guard cell number. The present results indicate that high flexibilities in stomatal density and guard cell size will change in response to water status, and this process may be closely associated with photosynthesis and water use efficiency.     

松嫩草地全叶马兰夏季与秋季光合及蒸腾作用的比较

在生长季晴天条件下,夏季和秋季松嫩草地全叶马兰的光合,蒸腾作用的日变化均为双峰曲线,但不同季节有所差异,夏季日均净光合速率与蒸腾速率均高于秋季,光合和蒸腾作用与环境因子的植物内部因子之间有密切关系。分析表明,叶片净光合速率与有效光辐射呈极显著相关,与气孔阻力,胞间CO2呈负相关;蒸腾速率与有效光辐射呈极显著相关,与叶温,饱和差呈正相关,与气孔阻力,胞间CO2浓度呈负相关,有效光辐射是影响光合和蒸腾作用诸因子中的主导因子,而气孔阻力变化则在调节光合和蒸腾中起着重要作用,蒸腾速率午降主要由于光辐射强,叶温高,湿度低,植物体缺水,气孔部分关闭所致。     

Decreased Rubisco activity during water stress is not induced by decreased relative water content but related to conditions of low stomatal conductance and chloroplast CO2 concentration

Rubisco activity decreases under water stress, for reasons as yet unclear. Here, the covariation of stomatal conductance (gs) and relative water content (RWC), often observed during water stress, was impaired to assess the separate effects of these factors on Rubisco activity. Three different treatments were applied to soybean (Glycine max) and tobacco (Nicotiana tabacum): leaf desiccation (LD), in which stomatal closure was accompanied by large decreases of RWC; water stress (WS), in which minor decreases of RWC were observed along with stomatal closure; and exogenous application of abscisic acid (ABA), which triggered stomatal closure without changing RWC. Decreased RWC did not induce decreased initial Rubisco activity, which was impaired only in soybean by 40% when the gs dropped below 50 mmol m(-2) s(-1), regardless of the treatment. The mechanism for decreased activity differed among treatments, owing to decreased activation in LD and to total activity and protein content in WS and ABA. Despite the occurrence of Rubisco regulation, CO2 availability in the chloroplast, not impairment of Rubisco activity, limits photosynthesis during WS.     

Xylem transport and shoot accumulation of lumichrome, a newly recognized rhizobial signal, alters root respiration, stomatal conductance, leaf transpiration and photosynthetic rates in legumes and cereals

* Root respiration, stomatal conductance, leaf transpiration and photosynthetic rates were measured in phytotron and field-grown plants following the application of 5 or 10 nM lumichrome, 10 nM ABA (abscisic acid) and 10 ml of 0.2 OD600 infective rhizobial cells. * Providing soybean and cowpea roots with their respective homologous rhizobia and/or purified lumichrome increased the concentration of this molecule in xylem sap and leaf extracts. Relative to control, rhizobial inoculation and lumichrome application significantly increased root respiration in maize, decreased it in lupin, but had no effect on the other test species. * Applying either lumichrome (10 nM), infective rhizobial cells or ABA to roots of plants for 44 h in growth chambers altered leaf stomatal conductance and transpiration in cowpea, lupin, soybean, Bambara groundnut and maize, but not in pea or sorghum. Where stomatal conductance was increased by lumichrome application or rhizobial inoculation, it resulted in increased leaf transpiration relative to control plants. Treating roots of field plants of cowpea with this metabolite up to 63 d after planting showed decreased stomatal conductance, which affected CO2 intake and reduction by Rubisco. * The effect of rhizobial inoculation closely mirrored that of lumichrome application to roots, indicating that rhizobial effects on these physiological activities were most likely due to lumichrome released into the rhizosphere.     

Proteomic response of rice seedling leaves to elevated CO2 levels

Previous investigations of plant responses to higher CO 2 levels were mostly based on physiological measurements and biochemical assays. In this study, a proteomic approach was employed to investigate plant response to higher CO 2 levels using rice as a model. Ten-day-old seedlings were progressively exposed to 760 ppm, 1140 ppm, and 1520 ppm CO 2 concentrations for 24 h each. The net photosynthesis rate ( P n), stomatal conductance ( G s), transpiration rate ( E), and intercellular to ambient CO 2 concentration ratio ( C i/ C a) were measured. P n, G s, and E showed a maximum increase at 1140 ppm CO 2, but further exposure to 1520 ppm for 24 h resulted in down regulation of these. Proteins extracted from leaves were subjected to 2-DE analysis, and 57 spots showing differential expression patterns, as detected by profile analysis, were identified by MALDI-TOF/TOF-MS. Most of the proteins belonged to photosynthesis, carbon metabolism, and energy pathways. Several molecular chaperones and ascorbate peroxidase were also found to respond to higher CO 2 levels. Concomitant with the down regulation of P n and G s, the levels of enzymes of the regeneration phase of the Calvin cycle were decreased. Correlations between the protein profiles and the photosynthetic measurements at the three CO 2 levels were explored.     

星星草耐盐碱生理机制再探讨

用不同浓度Na2CO3处理星星草幼苗,7d后测定叶片相对电导率、渗透势、光合速率（Pn）、气孔导度（Gs）、胞间CO2浓度（Ci）、蒸腾强度（Tr）、水分利用效率（WUE）等生理指标,以研究星星草抗盐碱的生理机制。结果表明,低浓度Na2CO3胁迫下,星星草幼苗可通过渗透调节来减轻细胞所受的伤害,因而膜透性受影响较小,相对电导率上升较少,而Gs基本保持不变,Pn、Tr还有所促进,WUE有所增加,这些结果说明,星星草幼苗对低浓度碱性盐具有一定抗性;但犀星草幼苗的这种抗性有一定限度,随着Na2CO3胁迫浓度的增加,Gs、Tr、Pn依次受到抑制,最终膜透性急剧增加,说明星星草幼苗已受损严重,这些变化具有较强的顺序性。     

干旱区胡杨光合作用对高温和CO2浓度的响应

采用LI-6400便携式光合作用测定仪实测的塔里木河下游胡杨(Populus euphratica oliv)光合作用参数,探讨了不同地下水埋深下的胡杨光合作用对CO2浓度增加和温度升高的响应.结果表明:(1)CO2浓度升高减小了胡杨气孔导度,促进了光合速率、胞间CO2浓度和水分利用效率的增加,但不同地下水埋深下,胡杨光合作用参数对CO2浓度升高的响应不同,干旱环境(地下水埋深较深)下的响应程度大于水分适宜(地下水埋深浅)环境下的响应;(2) 高温引起胡杨气孔发生不完全关闭,导致了光合作用的光抑制发生,从而降低了胡杨光合速率,但降低程度受水分条件的影响,地下水埋深较深环境下的影响程度大于地下水埋深浅的;(3)地下水埋深是控制干旱区胡杨光合作用对CO2浓度和温度升高的根本因素,6m是胡杨生长正常的临界地下水埋深,地下水埋深>6m,胡杨即遭到水分胁迫,地下水埋深>7m,胡杨即受到了较严重的水分胁迫.     

Growth characteristics and antioxidant metabolism of moongbean genotypes differing in photosynthetic capacity subjected to water deficit stress

Abstract

Two genotypes (Pusa 9531 and PS 16) of moongbean [Vigna radiata (L) Wilczek], differing in photosynthetic capacity were grown for 30 days in earthen pots at three field capacities (100, 75 and 50%), and the possible role of biochemical alterations and antioxidant metabolism in conferring photosynthetic capacity was determined by measuring Rubisco activity, photosynthetic traits, lipid peroxidation and assaying activities of the central components of antioxidant defence system. Growth, Rubisco activity, photosynthetic traits and soluble protein content decreased significantly with decreasing field capacity (FC) from 100 to 50%. Levels of TBARS, H₂O₂, electrolyte leakage and proline contents increased with decreasing FC. Activities of SOD and GR increased in both genotypes with decreasing FC; the CAT and APX activities over-expressed only at mild (75%) FC but not at severe (50%) FC. There were found genotype-dependent alterations in growth, photosynthetic traits, Rubisco activity and antioxidant metabolism when exposed to water deficit. Decline in efficiency of the H₂O₂-decomposing system at severe drought was responsible for oxidative damage occurring in both the genotypes. The differential responses of antioxidative enzymes in the two genotypes were the result of their ability to protect photosynthetic apparatus and alleviate water deficit stress.     

Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited

There is a long-standing controversy as to whether drought limits photosynthetic CO2 assimilation through stomatal closure or by metabolic impairment in C3 plants. Comparing results from different studies is difficult due to interspecific differences in the response of photosynthesis to leaf water potential and/or relative water content (RWC), the most commonly used parameters to assess the severity of drought. Therefore, we have used stomatal conductance (g) as a basis for comparison of metabolic processes in different studies. The logic is that, as there is a strong link between g and photosynthesis (perhaps co-regulation between them), so different relationships between RWC or water potential and photosynthetic rate and changes in metabolism in different species and studies may be 'normalized' by relating them to g. Re-analysing data from the literature using light-saturated g as a parameter indicative of water deficits in plants shows that there is good correspondence between the onset of drought-induced inhibition of different photosynthetic sub-processes and g. Contents of ribulose bisphosphate (RuBP) and adenosine triphosphate (ATP) decrease early in drought development, at still relatively high g (higher than 150 mmol H20 m(-2) s(-1)). This suggests that RuBP regeneration and ATP synthesis are impaired. Decreased photochemistry and Rubisco activity typically occur at lower g (<100 mmol H20 m(-2) s(-1)), whereas permanent photoinhibition is only occasional, occurring at very low g (<50 mmol H20 m(-2) s(-1)). Sub-stomatal CO2 concentration decreases as g becomes smaller, but increases again at small g. The analysis suggests that stomatal closure is the earliest response to drought and the dominant limitation to photosynthesis at mild to moderate drought. However, in parallel, progressive down-regulation or inhibition of metabolic processes leads to decreased RuBP content, which becomes the dominant limitation at severe drought, and thereby inhibits photosynthetic CO2 assimilation.     

Morphology and Stomatal Function of Douglas Fir Needles Exposed to Climate Change: Elevated CO2 and Temperature

Climate change may have an impact on the productivity of conifer trees by influencing the morphology (size and surface characteristics) and function (capacity for gas exchange) of conifer needles. In order to test the responses of needles to climatic variables, Douglas fir (Pseudotsuga menziesii [Mirb.] Franco), saplings were grown in sunlit controlled environment chambers at ambient or elevated (+200 parts per million above ambient) CO2 and at ambient or elevated temperature (+4 degrees C above ambient). Needle characteristics, including length, width, area, stomatal density (stomata per mm2), percentage of stomatal occlusion, and the morphology of epicuticular wax, were evaluated. Needle function was evaluated as stomatal conductance to water vapor and transpiration. Needle length increased significantly with elevated temperature but not with elevated CO2. Neither elevated CO2 nor elevated temperature affected stomatal density or stomatal number in these hypostomatous needles. Epicuticular wax was less finely granular at elevated than at ambient temperature and was similar in appearance at elevated and ambient CO2. Stomatal conductance and transpiration increased with elevated temperature and associated increased vapor pressure deficit; however, neither conductance nor transpiration was affected by elevated CO2. These results indicate that simulated climate change influences Douglas fir needle structure and function.     

Transpiration of spring barley under nitrogen and phosphorus deficiency during leaf wilting

In an open gas exchange system with a thermocouple psychrometer the transpiration rate of the first leaf in 8-day plants of spring barley was measured in dependence on the water saturation deficit (ΔW _sat). The plants were cultivated in Richter’s nutrient solution, either complete, or deficient in nitrogen or phosphorus. The cuticular transpiration (as measured in the dark) was unaffected by N and P deficiency. The N deficiency reduced the transpiration rate by increasing stomatal resistance since full water saturation of the leaf (67% rate of the control variant) up to stomatal closing at Δ W_sat = 14%. The P deficiency does not affect the transpiration rate at initial phases of wilting, but the stomata close only at a higher Δ W_sat (25%) than those in the control.     

Interactive effects of elevated CO2, warming, and drought on photosynthesis of Deschampsia flexuosa in a temperate heath ecosystem

Global change factors affect plant carbon uptake in concert. In order to investigate the response directions and potential interactive effects, and to understand the underlying mechanisms, multifactor experiments are needed. The focus of this study was on the photosynthetic response to elevated CO(2) [CO2; free air CO(2) enrichment (FACE)], drought (D; water-excluding curtains), and night-time warming (T; infrared-reflective curtains) in a temperate heath. A/C(i) curves were measured, allowing analysis of light-saturated net photosynthesis (P(n)), light- and CO(2)-saturated net photosynthesis (P(max)), stomatal conductance (g(s)), the maximal rate of Rubisco carboxylation (V(cmax)), and the maximal rate of ribulose bisphosphate (RuBP) regeneration (J(max)) along with leaf δ(13)C, and carbon and nitrogen concentration on a monthly basis in the grass Deschampsia flexuosa. Seasonal drought reduced P(n) via g(s), but severe (experimental) drought decreased P(n) via a reduction in photosynthetic capacity (P(max), J(max), and V(cmax)). The effects were completely reversed by rewetting and stimulated P(n) via photosynthetic capacity stimulation. Warming increased early and late season P(n) via higher P(max) and J(max). Elevated CO(2) did not decrease g(s), but stimulated P(n) via increased C(i). The T×CO2 synergistically increased plant carbon uptake via photosynthetic capacity up-regulation in early season and by better access to water after rewetting. The effects of the combination of drought and elevated CO(2) depended on soil water availability, with additive effects when the soil water content was low and D×CO2 synergistic stimulation of P(n) after rewetting. The photosynthetic responses appeared to be highly influenced by growth pattern. The grass has opportunistic water consumption, and a biphasic growth pattern allowing for leaf dieback at low soil water availability followed by rapid re-growth of active leaves when rewetted and possibly a large resource allocation capability mediated by the rhizome. This growth characteristic allowed for the photosynthetic capacity up-regulations that mediated the T×CO2 and D×CO2 synergistic effects on photosynthesis. These are clearly advantageous characteristics when exposed to climate changes. In conclusion, after 1 year of experimentation, the limitations by low soil water availability and stimulation in early and late season by warming clearly structure and interact with the photosynthetic response to elevated CO(2) in this grassland species.     

Regulation of Photosynthetic Rate of Two Sunflower Hybrids under Water Stress

The effect of short-term water stress on photosynthesis of two sunflower hybrids (Helianthus annuus L. cv Sungro-380 and cv SH-3622), differing in productivity under field conditions, was measured. The rate of CO₂ assimilation of young, mature leaves of SH-3622 under well-watered conditions was approximately 30% greater than that of Sungro-380 in bright light and elevated CO₂; the carboxylation efficiency was also larger. Growth at large photon flux increased assimilation rates of both hybrids. The changes in leaf composition, including cell numbers and sizes, chlorophyll content, and amounts of total soluble and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) protein, and in Rubisco activity and amount of ribulose-1,5-bisphosphate (RuBP) were determined to assess the factors regulating the differences in assimilation of the hybrids at high and low water potentials. The amounts of chlorophyll, soluble protein, Rubisco protein and the initial activity of Rubisco and its activation state did not differ significantly between hybrids. However, unstressed leaves of SH-3622 had more, smaller cells per unit area and 60% more RuBP per unit leaf area than that of Sungro-380. Water stress developing over 4 days decreased the assimilation of both hybrids similarly. Changes in the amounts of chlorophyll, soluble and Rubisco protein, and Rubisco activity and activation state were small and were not sufficient to explain the decrease in photosynthesis; neither was decreased stomatal conductance (or stomatal “patchiness”). Reduction of photosynthesis per unit leaf area from 25 to 5 micromoles CO₂ per square meter per second in both hybrids was caused by a decrease in the amount of RuBP from approximately 130 to 40 micromoles per square meter in SH-3622 and from 80 to 40 micromoles per square meter in Sungro. Differences between hybrids and their response to water stress is discussed in relation to control of RuBP regeneration.     

Stomatal regulation of photosynthesis in apple leaves: evidence for different water-use strategies between two cultivars

BACKGROUND AND AIMS: Leaf responses to environmental conditions have been frequently described in fruit trees, but differences among cultivars have received little attention. This study shows that parameters of Farquhar's photosynthesis and Jarvis' stomatal conductance models differed between two apple cultivars, and examines the consequences of these differences for leaf water use efficiency. METHODS: Leaf stomatal conductance (g(sw)), net CO2 assimilation rate (A(n)), respiration (R(d)) and transpiration (E) were measured during summer in 8-year-old 'Braeburn' and 'Fuji' apple trees under well-watered field conditions. Parameters of Farquhar's and Jarvis' models were estimated, evaluated and then compared between cultivars. Leaf carbon isotope discrimination (delta(13)C) was measured at the end of the growing season. KEY RESULTS: A single positive relationship was established between V(Cmax) (maximum carboxylation rate) and N(a) (leaf nitrogen concentration per unit area), and between J(max) (maximum light-driven electron transport rate) and N(a). A higher leaf R(d) was observed in 'Fuji'. The g(sw) responded similarly to increasing irradiance and leaf temperature in both cultivars. g(sw) responded to lower vapour pressure deficit in 'Fuji' than in 'Braeburn'. Maximal conductance (g(swmax)) was significantly smaller and A(n) was more limited by g(sw) in 'Braeburn' than 'Fuji'. Lower g(sw), E and higher intrinsic water use efficiency were shown in 'Braeburn' and confirmed by smaller leaf delta(13)C compared with 'Fuji' leaves. CONCLUSIONS: The use of functional model parameters allowed comparison of the two cultivars and provided evidence of different water use 'strategies': 'Braeburn' was more conservative in water use than 'Fuji', due to stomatal limitation of A(n), higher intrinsic water use efficiency and lower delta(13)C. These physiological traits need to be considered in relation to climate adaptation, breeding of new cultivars and horticultural practice.     

Interactions between water deficit, ABA, and provenances in Picea asperata

The effects of exogenous abscisic acid (ABA) on the acclimation of Picea asperata to water deficit were investigated in two populations originating from wet and dry climate regions of China. Exogenous ABA was sprayed onto the leaves, and changes in plant growth and structure, gas exchange, water use efficiency (WUE), endogenous ABA content, and antioxidant enzyme levels were monitored. The results demonstrated that ABA application affected the two P. asperata populations in different ways during the water deficit. ABA application resulted in significantly lower CO(2) assimilation rates (A) under water deficit in plants from the wet climate population, whereas there were no significant changes in this parameter in the dry climate population. On the other hand, ABA application significantly decreased the dry shoot biomass, stomatal conductance (g(s)), transpiration rate (E), and malondialdehyde (MDA) content, and it significantly increased the leaf mass per area (LMA), root/shoot ratio (Rs), fine root/total root ratio (Ft), WUE, ABA content, and the superoxide dismutase (SOD), ascorbate peroxidase (APX), and catalase (CAT) activities under water-deficit conditions in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet climate population. The results clearly demonstrated that sensitivity to an exogenous ABA application is population-dependent in P. asperata. Direct evidence is presented that variation in physiological mechanisms rather than different rates of ABA absorption explain the population differentiation in the sensitivity to exogenous ABA, and that the physiological basis for the amplified response to water deficit caused by exogenous ABA, present mainly in the dry climate population, is related to internal ABA accumulation. These results provide evidence for adaptive differentiation between populations of P. asperata, and they support the expected relationship between environmental heterogeneity and the magnitude of plastic responses in plant populations.     

Chloroplast downsizing under nitrate nutrition restrained mesophyll conductance and photosynthesis in rice (Oryza sativa L.) under drought conditions

The phenomenon whereby ammonium enhances the tolerance of rice seedlings (Oryza sativa L., cv. 'Shanyou 63' hybrid indica China) to water stress has been reported in previous studies. To study the intrinsic mechanism of biomass synthesis related to photosynthesis, hydroponic experiments supplying different nitrogen (N) forms were conducted; water stress was simulated by the addition of polyethylene glycol. Water stress decreased leaf water potential (Ψ(leaf)) under nitrate nutrition, while it had no negative effect under ammonium nutrition. The decreased Ψ(leaf) under nitrate nutrition resulted in chloroplast downsizing and subsequently decreased mesophyll conductance to CO(2) (g(m)). The decreased g(m) and stomatal conductance (g(s)) under nitrate nutrition with water stress restrained the CO(2) supply to the chloroplast and Rubisco. The relatively higher distribution of leaf N to Rubisco under ammonium nutrition might also be of benefit for photosynthesis under water stress. In conclusion, chloroplast downsizing induced a decline in g(m), a relatively higher decrease in g(s) under nitrate nutrition with water stress, restrained the CO(2) supply to Rubisco and finally decreased the photosynthetic rate.     

贝加尔针茅不同枝条叶片蒸腾特性与水分利用效率对瞬时CO_2和光照变化的响应

利用人工光源测量了不同 CO2 浓度条件下贝加尔针茅 (Stipa bacailensis)营养枝条与生殖枝条叶片的净光合速率 (PN)、蒸腾速率 (E)、气孔导度 (gs)、胞间 CO2 浓度 (Ci)及叶面饱和蒸气压亏缺 (VPD)。营养枝与生殖枝 PN 及 E均随 CO2 浓度升高而增大 ,但 PN 增加幅度较大 ,E增加幅度较小。在高 CO2 浓度 (14 0 0 μmol/m ol)条件下 ,营养枝叶片最大 PN(2 7.2 3μmol CO2 /(m2 · s) )大于生殖枝 (17.13μm ol CO2 /(m2 · s) )。营养枝与生殖枝之间 E呈极显著差异。营养枝与生殖枝水分利用效率 (WUE= PN/E)均随 CO2 浓度升高而增大 ,生殖枝 WUE略高于营养枝 ,但差异未达到显著水平。光合速率的显著增加是贝加尔针茅水分利用效率随 CO2 浓度升高而增加的主要影响因素。CO2 浓度相对稳定条件下 (35 0 μmol/mol) ,生殖枝与营养枝 PN 与 E均随模拟光辐射 (SPR)强度增加而增大 ,但增幅逐渐趋缓 ,营养枝最大 PN 及 E均大于生殖枝。当 SPR强度从 0增加到 4 0 0 μmol/(m2 · s)过程中 ,营养枝与生殖枝叶片水分利用效率均呈陡然增大趋势 ,随着 SPR的进一步增强 ,WUE缓慢增大并在较高值附近达到波动平衡。贝加尔针茅营养枝与生殖枝之间的 gs差异是 PN 与 E差异的主要影响因素 ,也决定了 WUE对 CO2 浓度和模拟光辐