Changes in Stomatal Conductance,Transpiration and Water Use Efficiency of Ten Species Experienced in High CO2 Concentrations in Biosphere 2

The stomotal conductance, transpiration and water use efficiency (WUE) were measured using a LI-6400 portable photosynthesis system for 5 tropical rain forest species and 5 desert species in Biosphere 2, USA. All the species have experienced in very high CO2 ( > 2 200 μmol• mol- 1 ) for more than 4.5 years. The results showed that the stomatal conductance and transpiration of rain forest species decreased from ( 127.4 ± 65.6) and (2.04 ± 0.61 ) mmol• m- 2•s- 1 to (61.3 + 30.5) and ( 1.54 ± 0.65 ) mmol• m-2• s -1 respectively, while WUE increased from (2.90 ± 0.55) to (8.45 ± 2.71) μmol CO2 •mmo1-1 H2O, with CO2 increasing from 350 – 400 to 700 – 820 μmol• mol-l. For the desert species, stomatal conductance and transpiration decreased from respectively (142.8±94.6) and (2.09±0.71) mmol•m-2•s-1 to (57.7±35.8) and (1.36±0.52) mmolm-2•s-l, but WUE increased from (4.69 ± 1.39) to (9.68 ± 1.61) μmol CO2•mmo1-1 H2O, with the CO2 increase from 320 - 400 to 820 – 850 μtmol• mol- 1. The stomatal conductance, transpiration and WUE were less influenced by light intensity under high CO2 than low CO2 concentrations. Most rain forest species reached their light saturation points at light intensity of 500 μmol• m-2•s-1, while desert species at 1 000 μmol•m-2•s-1. Among different species, the desert C3 tree, Nicotiana glauca Grah., had the highest decrease in stomatal conductance and transpiration and the highest increase in WUE, by 78%, 69% and 310% respectively. The enhancement of increasing CO2 to the stomatal, transpiration and WUE of species with different photosynthesis pathway and life forms in Biosphere 2 could be concluded as: C3 species > C4 species, and desert C3 species > rain forest C3 species.     

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.     

铁皮石斛叶片光合作用的碳代谢途径

 利用LI-6400光合测定系统测定了不同天气条件下铁皮石斛（Dendrobium officinale）叶片24h CO2吸收的动态以及CO2吸收对光强和温度的响应。晴天的白天和夜间铁皮石斛都能吸收CO2,中午CO2吸收速率为负值, CO2的交换方式具景天酸代谢途径(CAM)的特点。阴雨天,只有白天吸收CO2,夜间表现为暗呼吸,光合作用碳代谢的途径为C3途径。在多云的天气条件下,白天吸收CO2,并持续至日落后。夜间21∶00仍有CO2吸收,23∶00以后至次日凌晨处于暗呼吸状态。在500 μmol·m-2·s-1光照件下,20℃出现最大CO2吸收值。在夜间,25℃时CO2的吸收速率最高。有光和无光条件下,低温或高温引起CO2吸收速率下降均为非气孔因素所致。晴天上午,铁皮石斛叶片的表观量子产额为0.035,光合补偿点为2.9μmol·m-2·s-1,饱和光强为500μmol·m-2·s-1,强光下出现光抑制现象。叶片受到强光预先照射后,即使光照减弱光抑制效应仍保持一段时间,致使光合补偿点升高,表观量子产额下降,相同光强下的CO2吸收效率降低。结果表明：铁皮石斛为兼性CAM植物,随着环境条件的变化,其光合作用在景天酸代谢途径(CAM)与C3途径间变化。     

全球变化条件下植物个体的生理生态学模型

天气模型中应用随机模拟方法,产生以天或小时为时间间隔的气温、降水、相对湿度、云量、太阳辐射等天气要素的动态变化时间序列。利用北京地区近30年天气资料进行了模拟验证,模拟结果与实际的天气变化进程相符。生理生态模型描述了净光合速率、气孔传导度、蒸腾速率、水分利用效率的变化机理。结合开顶式CO_2浓度倍增大豆(Glycine max(L.)Merr.)生长实验,分析了这些生理生态特性在全球变化下的动态响应机制,并进行了模拟预测。结果表明:CO_2浓度倍增情况下,净光合速率提高45％,其中光量子效率显著增加,而CO_2传导系数略有下降;气孔传导度、蒸腾速率下降约30％;水分利用效率随CO_2浓度增加几乎呈线性增长,倍增后提高近一倍。     

小麦和大豆叶片的气孔不均匀关闭现象

用14CO2放射自显影的方法研究了田间小麦和大豆叶片在水分胁迫下的气孔关闭状况。正常浇水的小麦和大豆叶片呈现出对14CO2的均匀吸收。在小麦与大豆"片水势分别降至-1．75和-1．32MPa的土壤干旱条件下,两种作物叶片都发生气孔不均匀关闭。离休叶片在空气中快速脱水易引起气孔不均匀关闭。正常供水小麦叶片在晴天中午明显的光合午休时,无CO2的不均匀吸收。某些晴天中午,在大豆光合午休低谷时段观察到较明显的气孔不均匀关闭,用气体交换资料计算出的细胞间隙CO2浓度并不随气孔年度的降低而下降,反而略有回升。     

Inhibition of stomatal opening in sunflower leaves by carbon monoxide,and reversal of inhibition by light

When leaves of Helianthus annuus, whose stomates had been opened in the dark in the absence of CO₂, were exposed to 25% carbon monoxide (CO), stomatal conductivity for water vapor decreased from about 0.4 to 0.2 cm·s^-1. The CO effect on stomatal aperture required a CO/O₂ ratio of about 25. As this ratio was decreased the stomata opened, indicating that inhibitio of cytochrome-c oxidase by CO is competitive in respect to O₂. Photosynthetically active red light was unable to reverse CO-induced stomatal closure even at high irradiances, when CO₂ was absent. When it was present, stomatal opening was occasionally, but not consistently observed. Carbon monoxide did not inhibit photosynthetic carbon reduction in leaves of Helianthus.In contrast to red light, very weak blue light (405 nm) increased the stomatal aperture in the presence of CO. It also increased leaf ATP/ADP ratios which had been decreased in the presence of CO. The blue-light effect was not related to photosynthesis. Neither could it be explained by photodissociation of the cytochrome a ₃-CO complex which has an absorption maximum at 430 nm. The data indicate that ATP derived from mitochondrial oxidative phosphorylation provides energy for stomatal opening in sunflower leaves in the dark as well as in the light. Indirect transfer of ATP from chloroplasts to the cytosol via the triose phosphate/phosphoglycerate exchange which is mediated by the phosphate translocator of the chloroplast envelope can support stomatal opening only if metabolite concentrations are high enough for efficient shuttle transfer of ATP. Blue light causes stomatal opening in the presence of CO by stimulating ATP synthesis.     

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.     

CO2浓度升高对干旱胁迫下小麦光合作用和抗氧化酶活性的影响

在CO     

Limitations to photosynthesis of lettuce grown under tropical conditions: alleviation by root-zone cooling.

Aerial parts of lettuce plants were grown under natural tropical fluctuating ambient temperatures, but with their roots exposed to two different root-zone temperatures (RZTs): a constant 20 degrees C-RZT and a fluctuating ambient (A-) RZT from 23-40 degrees C. Plants grown at A-RZT showed lower photosynthetic CO2 assimilation (A), stomatal conductance (gs), midday leaf relative water content (RWC), and chlorophyll fluorescence ratio Fv/Fm than 20 degrees C-RZT plants on both sunny and cloudy days. Substantial midday depression of A and g(s) occurred on both sunny and cloudy days in both RZT treatments, although Fv/Fm did not vary diurnally on cloudy days. Reciprocal temperature transfer experiments investigated the occurrence and possible causes of stomatal and non-stomatal limitations of photosynthesis. For both temperature transfers, light-saturated stomatal conductance (gs sat) and photosynthetic CO2 assimilation (A(sat)) were highly correlated with each other and with midday RWC, suggesting that A was limited by water stress-mediated stomatal closure. However, prolonged growth at A-RZT reduced light- and CO2-saturated photosynthetic O2 evolution (Pmax), indicating non-stomatal limitation of photosynthesis. Tight temporal coupling of leaf nitrogen content and P(max) during both temperature transfers suggested that decreased nutrient status caused this non-stomatal limitation of photosynthesis.     

Responses to Doubled CO2 in Medicago sativa: Studies on Ecophysiology and Simulation Modelling

A world-wide spread forage grass, Medicago sativa, was grown in two open-top chambers maintained at either normal (350 μmol · mol-1) or doubled (700 μmol · mol-1) CO2 concentration, from seedling to maturity. During the whole growth season, ecophysiological measurements and observations were conducted over different phenological stages and the main results were as follows: (1) With similar environmental factors, in terms of RH (relative humidity), irradiance, and watering, a slight shift in temperature (about 0.77℃, averaged over the whole growth season) within the chamber maintained at doubled CO2 did not affect instantaneous physiological processes at leaf level, but had some impacts on the effect of doubled CO2 over a long period. (2) Over the whole growth season, physiological variables showed differences between two chambers. The net photosynthesis of plant grown under higher CO2 increased by 18.7 %, while stomatal conductance fell slightly as compared with that of the control. So was the water use efficiency which was also 30.1% higher than the control. Based on the above results from field studies, we concluded that photosynthetically active radiation (PAR) and RH were the main factors affecting photosynthesis and stomatal behavior. Then we combined a widely accepted model of C3 leaf photosynthesis with an empirical model of stomatal conductance and made some modifications according to our experiments. This model was parameterized using field data sets of net CO2 assimilation, stomatal conductance, intercellular CO2 concentration of plants grown at both doubled and control CO2 levels. Variances of main parameters between two treatments reflected some biochemical changes in leaf cell. The maximum efficiency of light energy conversion (α) increased by 22 % and light-saturated rate of electron transport (Jmax) rose by 15 %. The maximum stomatal conductance was slightly reduced by 8 %. The increases in parameters (α and Jmax) indicate accelerated biochemical processes in leaf cell, which means that the photosynthetic capacity of M. sativa may increase at elevated CO2. These results agree well with biochemical measurements at cell level.     

Physiological responses of beech and sessile oak in a natural mixed stand during a dry summer

Responses of CO2 assimilation and stomatal conductance to decreasing leaf water potential, and to environmental factors, were analysed in a mixed natural stand of sessile oak (Quercus petraea ssp. medwediewii) and beech (Fagus svlvatica L.) in Greece during the exceptionally dry summer of 1998. Seasonal courses of leaf water potential were similar for both species, whereas mean net photosynthesis and stomatal conductance were always higher in sessile oak than in beech. The relationship between net photosynthesis and stomatal conductance was strong for both species. Sessile oak had high rates of photosynthesis even under very low leaf water potentials and high air temperatures, whereas the photosynthetic rate of beech decreased at low water potentials. Diurnal patterns were similar in both species but sessile oak had higher rates of CO2 assimilation than beech. Our results indicate that sessile oak is more tolerant of drought than beech, due, in part, to its maintenance of photosynthesis at low water potential.     

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.     

Adaxial/abaxial specification in the regulation of photosynthesis and stomatal opening with respect to light orientation and growth with CO2 enrichment in the C4 species Paspalum dilatatum.

Whole-plant morphology, leaf structure and composition were studied together with the effects of light orientation on the dorso-ventral regulation of photosynthesis and stomatal conductance in Paspalum dilatatum cv. Raki plants grown for 6 wk at either 350 or 700 microl l(-1) CO(2). Plant biomass was doubled as a result of growth at high CO(2) and the shoot:root ratio was decreased. Stomatal density was increased in the leaves of the high CO(2)-grown plants, which had greater numbers of smaller stomata and more epidermal cells on the abaxial surface. An asymmetric surface-specific regulation of photosynthesis and stomatal conductance was observed with respect to light orientation. This was not caused by dorso-ventral variations in leaf structure, the distribution of phosphoenolpyruvate carboxylase (PEPC) and ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) proteins or light absorptance, transmittance or reflectance. Adaxial/abaxial specification in the regulation of photosynthesis results from differential sensitivity of stomatal opening to light orientation and fixed gradients of enzyme activation across the leaf.     

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.     

The effects of O3 fumigation during leaf development on photosynthesis of wheat and pea: An in vivo analysis

Previous studies have shown that short exposure of plants to high doses of ozone decreases subsequent photosynthesis; initially by reducing carboxylation capacity. This study tests the hypothesis that this is also the primary cause of loss of photosynthetic capacity in leaves affected by development under a low level of ozone. Triticum aestivum and Pisum sativum plants were exposed from germination to ozone in air (80 nmol mol^-1 for 7 hours per day, for 18 days. Leaves that had completed lamina expansion at this time were free of visible injury and light absorptance was unaffected. However, some significant changes in photosynthetic gas exchange were evident. Photosynthetic CO2 uptake at light saturation was decreased significantly by 35% in T. aestivum but was unchanged in P. sativum. The reduction in photosynthesis of T. aestivum was accompanied by a 31% decline in the maximum velocity of carboxylation measured in vivo. Decreased stomatal conductance did not contribute to this reduction of photosynthesis because there was no significant change in the stomatal limitation to CO2. Processes directly dependent upon photochemical reactions; that is, the quantum yield of CO2 uptake and capacity for regeneration of ribulose 1,5-bisphosphate were not affected by O3 fumigation in either species. This suggests that for wheat, the quantitative cause of decreased photosynthetic rate in vivo is a decrease in the quantity of active ribulose-1,5- bisphosphate carboxylase-oxygenase.     

高温胁迫对新疆榛光合参数和叶绿素荧光特性的影响

在5个温度梯度处理下,研究高温胁迫对4种新疆榛光合参数和叶绿素荧光特性的影响.结果表明:随着温度从25℃持续升高至45℃,新疆榛叶片的净光合速率、气孔导度、胞间CO2浓度、水分利用效率和光能利用效率逐渐降低,且在35～ 45℃之间降幅最大;光系统Ⅱ的实际光化学效率、电子传递速率和光化学猝灭系数随温度的升高缓慢上升,至35℃后急速下降;蒸腾耗水和热耗散随温度的升高而增大.4种新疆榛品种中,新榛3号的光合作用对高温的耐受力较高,属耐热性品种.     

根际CO2浓度升高对网纹甜瓜光合特性及产量和品质的影响

采用雾培植株根际通CO₂处理方式,研究了开花结果期根际CO₂浓度升高对网纹甜瓜光合作用及产量和品质的影响.结果表明：在网纹甜瓜果实发育期间,与350 μL·L^-1（对照）处理相比,根际2500和5000 μL CO₂·L^-1处理的叶片光合色素含量、净光合速率（P_n）、气孔导度（g_s）、胞间CO₂浓度（C_i）及PSⅡ最大光化学效率（F_v/F_m）均不同程度降低,而气孔限制值（L_s）显著提高,且5000 μL CO₂·L^-1处理的变化幅度高于2500 μL CO₂·L^-1处理;单株产量、果实中维生素C和可溶性糖含量显著降低,有机酸含量显著提高.可见,网纹甜瓜果实发育期间根际CO₂浓度超过2500 μL·L^-1时,其光合作用及果实发育会受到显著抑制,从而导致产量和品质降低.
     

Recovery responses of photosynthesis,transpiration, and stomatal conductance in kidney bean following drought stress

Photosynthesis, transpiration, stomatal conductance and chlorophyll fluorescence characteristics were examined in kidney bean plants, with developing gradually water stress for several days after watering and then permitted to recover by re-watering. The photosynthetic rate, transpiration rate, and stomatal conductance decreased rapidly by withholding water for 2 days. The F_v/F_m of chlorophyll fluorescence characteristics slightly decreased when the water was withheld for 7 days. After re-watering the rate of recovery of photosynthesis, transpiration, and stomatal conductance decreased gradually as the days without watering became longer. The differences existed in rates of recovery of photosynthesis, transpiration, and stomatal conductance following drought stress. Among the fractional recoveries the highest was photosynthesis, and the lowest was stomatal conductance. Photosynthesis rate following drought stress was rapidly recovered until 2 days after re-watering, then recovered slowly. The critical time for the recovery of photosynthesis was recognized. The results show clearly a close correlation between the leaf water potential and the recovery level and speed of photosynthesis, transpiration, and stomatal conductance.     

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.     

The stomata of the fern Adiantum capillus-veneris do not respond to CO2 in the dark and open by photosynthesis in guard cells

The stomata of the fern Adiantum capillus-veneris lack a blue light-specific opening response but open in response to red light. We investigated this light response of Adiantum stomata and found that the light wavelength dependence of stomatal opening matched that of photosynthesis. The simultaneous application of red (2 micromol m(-2) s(-1)) and far-red (50 micromol m(-2) s(-1)) light synergistically induced stomatal opening, but application of only one of these wavelengths was ineffective. Adiantum stomata did not respond to CO2 in the dark; the stomata neither opened under a low intercellular CO2 concentration nor closed under high intercellular CO2 concentration. Stomata in Arabidopsis (Arabidopsis thaliana), which were used as a control, showed clear sensitivity to CO2. In Adiantum, stomatal conductance showed much higher light sensitivity when the light was applied to the lower leaf surface, where stomata exist, than when it was applied to the upper surface. This suggests that guard cells likely sensed the light required for stomatal opening. In the epidermal fragments, red light induced both stomatal opening and K+ accumulation in guard cells, and both of these responses were inhibited by a photosynthetic inhibitor, 3-(3,4-dichlorophenyl)-1,1-dimethylurea. The stomatal opening was completely inhibited by CsCl, a K+ channel blocker. In intact fern leaves, red light-induced stomatal opening was also suppressed by 3-(3,4-dichlorophenyl)-1,1-dimethylurea. These results indicate that Adiantum stomata lack sensitivity to CO2 in the dark and that stomatal opening is driven by photosynthetic electron transport in guard cell chloroplasts, probably via K+ uptake.