Long-term growth of soybean at elevated [CO2] does not cause acclimation of stomatal conductance under fully open-air conditions

Accurately predicting plant function and global biogeochemical cycles later in this century will be complicated if stomatal conductance (g(s)) acclimates to growth at elevated [CO(2)], in the sense of a long-term alteration of the response of g(s) to [CO(2)], humidity (h) and/or photosynthetic rate (A). If so, photosynthetic and stomatal models will require parameterization at each growth [CO(2)] of interest. Photosynthetic acclimation to long-term growth at elevated [CO(2)] occurs frequently. Acclimation of g(s) has rarely been examined, even though stomatal density commonly changes with growth [CO(2)]. Soybean was grown under field conditions at ambient [CO(2)] (378 micromol mol(-1)) and elevated [CO(2)] (552 micromol mol(-1)) using free-air [CO(2)] enrichment (FACE). This study tested for stomatal acclimation by parameterizing and validating the widely used Ball et al. model (1987, Progress in Photosynthesis Research, vol IV, 221-224) with measurements of leaf gas exchange. The dependence of g(s) on A, h and [CO(2)] at the leaf surface was unaltered by long-term growth at elevated [CO(2)]. This suggests that the commonly observed decrease in g(s) under elevated [CO(2)] is due entirely to the direct instantaneous effect of [CO(2)] on g(s) and that there is no longer-term acclimation of g(s) independent of photosynthetic acclimation. The model accurately predicted g(s) for soybean growing under ambient and elevated [CO(2)] in the field. Model parameters under ambient and elevated [CO(2)] were indistinguishable, demonstrating that stomatal function under ambient and elevated [CO(2)] could be modelled without the need for parameterization at each growth [CO(2)].     

Heritable variation in stomatal responses to elevated CO2 in wild radish, Raphanus raphanistrum (Brassicaceae)

Rising atmospheric carbon dioxide may affect plant populations in the short term through effects on photosynthesis and carbon allocation, and over the long term as an agent of natural selection. To test for heritable effects of elevated CO2 on stomatal responses and plant fecundity in Raphanus raphanistrum, we grew plants from 12 paternal families in outdoor open-top chambers at ambient (35 Pa) or elevated (67 Pa) CO2. Contrary to results from a previous study of this species, total flower and fruit production were marginally lower under elevated CO2. Across families, stomatal index and guard cell length showed little response to CO2 enrichment, but these characters varied significantly among paternal families in both the direction and magnitude of their response to changing CO2. Although these family-by-CO2 interactions suggest that natural selection might affect stomatal characters when ambient CO2 levels increase, we found no significant correlation between either character and flower or fruit production. Therefore, our data suggest that while heritable variation for stomatal index and guard cell length exists in this population, selection due to increasing CO2 is not likely to act on these traits because they had no detectable effect on lifetime fecundity.     

高浓度二氧化碳对植物影响的研究进展

工业革命后全球大气CO2浓度持续上升,不仅对全球气候的变迁产生重大影响,而且对植物的形态、水分利用、蛋白质合成、光合、抗性、生长及生物量等都有不同程度的影响。高浓度CO2促进植物根、幼苗的生长,叶片增厚,降低气孔密度、气孔导度及蒸腾速率,增加水分利用效率、作物的产量及生物量,促进乙烯生物合成,增强植物的抗氧化能力。不同光合途径(C3、C4及CAM)及不同植被类型的植物对高浓度CO2的响应不同。长期和短期的高浓度CO2处理,植物响应方式有很大的差异,如短期高CO2处理使光合能力增强,而长期处理则使光合能力下调。     

植物对开放式CO2 浓度增高(FACE)的响应与适应研究进展

开放式CO2浓度增高(FACE)系统是近年研究植物对高CO2浓度响应和适应的新手段,它比以往密闭和半密闭系统对实验植物生长环境的干扰少.利用FACE系统进行研究更有助于正确地预测未来大气CO2浓度增高对植物的影响.该文结合作者的研究工作简要评介了FACE系统与以往密闭和半密闭式CO2浓度增高实验系统的不同之处以及近年来利用FACE系统所作的最新研究进展.     

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.     

盐胁迫对星星草幼苗光合特性的影响

对松嫩碱化草地上人工种植生长一至三年的星星草和自然生长的天然星星草的光合蒸腾特性的季节变化进行了测定。星星草光合速率的季节变化是单峰曲线,抽穗期光合速率最大。星星草的蒸腾速率则随生育期进程逐渐减小,但单位面积草地上星星草群体蒸腾速率的季节变化为单峰曲线,一年生、二年生和三年生星星草的最大值在开花期,而天然星星草则在抽穗期。星星草水分利用效率的季节变化也是单峰曲线,峰值均在抽穗期。     

开放式空气CO2浓度升高与作物/杂草的竞争关系

CO2浓度升高对植物的光合作用、呼吸作用和水分利用等生理过程产生直接影响,进而影响植物的生长繁殖,CO2浓度升高对于具有C3光合途径的植物较具C4光合途径的植物更为有益,由于许多重要的杂草是C4植物,而许多重要的作用是C3植物,CO2浓度升高对杂草/作物的相互关系将有重要影响,本文就全球CO2浓度升高和气候变化对杂草/作物之间竞争关系影响进行综述,同时针对目前研究现状和可持续农业的需要,提出CO2学浓度升高条件下杂草/作物之间竞争关系及未来农田杂草治理方面理论与实践中有待解决的问题。     

Simulation of the stomatal conductance of winter wheat in response to light, temperature and CO2 changes

BACKGROUND AND AIMS: The stomata are a key channel of the water cycle in ecosystems, and are constrained by both physiological and environmental elements. The aim of this study was to parameterize stomatal conductance by extending a previous empirical model and a revised Ball-Berry model. METHODS: Light and CO(2) responses of stomatal conductance and photosynthesis of winter wheat in the North China Plain were investigated under ambient and free-air CO(2) enrichment conditions. The photosynthetic photon flux density and CO(2) concentration ranged from 0 to 2000 micro mol m(-2) s(-1) and from 0 to 1400 micro mol mol(-1), respectively. The model was validated with data from a light, temperature and CO(2) response experiment. RESULTS: By using previously published hyperbolic equations of photosynthetic responses to light and CO(2), the number of parameters in the model was reduced. These response curves were observed diurnally with large variations of temperature and vapour pressure deficit. The model interpreted stomatal response under wide variations in environmental factors. CONCLUSIONS: Most of the model parameters, such as initial photon efficiency and maximum photosynthetic rate (P(max)), have physiological meanings. The model can be expanded to include influences of other physiological elements, such as leaf ageing and nutrient conditions, especially leaf nitrogen content.     

Optimal nitrogen allocation controls tree responses to elevated CO2

Despite the abundance of experimental data, understanding of forest responses to elevated CO2 is limited. Here I show that a key to previously unexplained production and leaf area responses lies in the interplay between whole-plant nitrogen (N) allocation and leaf photosynthesis. A simple tree growth model, controlled by net growth maximization through optimization of leaf area index (LAI) and plant N, is used to analyse CO2 responses in both young, expanding and closed, steady-state canopies. The responses are sensitive to only two independent parameters, the photosynthetic capacity per leaf N (a) and the fine-root N:leaf N ratio. The model explains observed CO2 responses of photosynthesis, production and LAI in four forest free air CO2 enrichment (FACE) experiments. Insensitivity of LAI except at low LAI, increase in light-use efficiency, and photosynthetic down-regulation (as a result of reduced leaf N per area) at elevated CO2 are all explained through the combined effects on a and leaf quantum efficiency. The model bridges the gap between the understanding of leaf-level and plant-level responses and provides a transparent framework for interpreting and linking structural (LAI) and functional (net primary production (NPP):gross primary production (GPP) ratio, light-use efficiency, photosynthetic down-regulation) responses to elevated CO2.     

大气CO2浓度和温度升高对作物生理生态的影响

论述了大气CO2浓度和温度升高下的植物生长,光合作用,产量以及水分养分利用效率等方面的研究进展,未来高CO2浓度下,光合作用速率有不同程度的提高,生物量和产量增加;气孔导度降低,水分利用效率（WUE）提高,一般地上部分和根系尤其是细根生物量增加,凋落物量随之增加,C/N比率提高,植物残体的腐解速率降低,CO2浓度升高后,会给根际微生物带来更多的底物,从而提高了微生物活性,加速养分的矿化过程,改善植物的养分状况。     

Changes in stomatal function and water use efficiency in potato plants with altered sucrolytic activity

As water availability for agriculture decreases, breeding or engineering of crops with improved water use efficiency (WUE) will be necessary. As stomata are responsible for controlling gas exchange across the plant epidermis, metabolic processes influencing solute accumulation in guard cells are potential targets for engineering. In addition to its role as an osmoticum, sucrose breakdown may be required for synthesis of other osmotica or generation of the ATP needed for solute uptake. Thus, alterations in partitioning of sucrose between storage and breakdown may affect stomatal function. In agreement with this hypothesis, potato (Solanum tuberosum) plants expressing an antisense construct targeted against sucrose synthase 3 (SuSy3) exhibited decreased stomatal conductance, a slight reduction in CO(2) fixation and increased WUE. Conversely, plants with increased guard cell acid invertase activity caused by the introduction of the SUC2 gene from yeast had increased stomatal conductance, increased CO(2) fixation and decreased WUE. (14)CO(2) feeding experiments indicated that these effects cannot be attributed to alterations in photosynthetic capacity, and most likely reflect alterations in stomatal function. These results highlight the important role that sucrose breakdown may play in guard cell function and indicate the feasibility of manipulating plant WUE through engineering of guard cell sucrose metabolism.     

Diffusive and metabolic limitations to photosynthesis under drought and salinity in C(3) plants

Drought and salinity are two widespread environmental conditions leading to low water availability for plants. Low water availability is considered the main environmental factor limiting photosynthesis and, consequently, plant growth and yield worldwide. There has been a long-standing controversy as to whether drought and salt stresses mainly limit photosynthesis through diffusive resistances or by metabolic impairment. Reviewing in vitro and in vivo measurements, it is concluded that salt and drought stress predominantly affect diffusion of CO(2) in the leaves through a decrease of stomatal and mesophyll conductances, but not the biochemical capacity to assimilate CO(2), at mild to rather severe stress levels. The general failure of metabolism observed at more severe stress suggests the occurrence of secondary oxidative stresses, particularly under high-light conditions. Estimates of photosynthetic limitations based on the photosynthetic response to intercellular CO(2) may lead to artefactual conclusions, even if patchy stomatal closure and the relative increase of cuticular conductance are taken into account, as decreasing mesophyll conductance can cause the CO(2) concentration in chloroplasts of stressed leaves to be considerably lower than the intercellular CO(2) concentration. Measurements based on the photosynthetic response to chloroplast CO(2) often confirm that the photosynthetic capacity is preserved but photosynthesis is limited by diffusive resistances in drought and salt-stressed leaves.     

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].     

Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2

To model the effect of increasing atmospheric CO2 on semi-arid grasslands, the gas exchange responses of leaves to seasonal changes in soil water, and how they are modified by CO2, must be understood for C3 and C4 species that grow in the same area. In this study, open-top chambers were used to investigate the photosynthetic and stomatal responses of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) grown at 360 (ambient CO2) and 720 micro mol mol-1 CO2 (elevated CO2) in a semi-arid shortgrass steppe. Assimilation rate (A) and stomatal conductance (gs) at the treatment CO2 concentrations and at a range of intercellular CO2 concentrations and leaf water potentials (psileaf) were measured over 4 years with variable soil water content caused by season and CO2 treatment. Carboxylation efficiency of ribulose bisphosphate carboxylase/oxygenase (Vc,max), and ribulose bisphosphate regeneration capacity (Jmax) were reduced in P. smithii grown in elevated CO2, to the degree that A was similar in elevated and ambient CO2 (when soil moisture was adequate). Photosynthetic capacity was not reduced in B. gracilis under elevated CO2, but A was nearly saturated at ambient CO2. There were no stomatal adaptations independent of photosynthetic acclimation. Although photosynthetic capacity was reduced in P. smithii growing in elevated CO2, reduced gs and transpiration improved soil water content and psileaf in the elevated CO2 chambers, thereby improving A of both species during dry periods. These results suggest that photosynthetic responses of C3 and C4 grasses in this semi-arid ecosystem will be driven primarily by the effect of elevated CO2 on plant and soil water relations.     

What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2

Free-air CO(2) enrichment (FACE) experiments allow study of the effects of elevated [CO(2)] on plants and ecosystems grown under natural conditions without enclosure. Data from 120 primary, peer-reviewed articles describing physiology and production in the 12 large-scale FACE experiments (475-600 ppm) were collected and summarized using meta-analytic techniques. The results confirm some results from previous chamber experiments: light-saturated carbon uptake, diurnal C assimilation, growth and above-ground production increased, while specific leaf area and stomatal conductance decreased in elevated [CO(2)]. There were differences in FACE. Trees were more responsive than herbaceous species to elevated [CO(2)]. Grain crop yields increased far less than anticipated from prior enclosure studies. The broad direction of change in photosynthesis and production in elevated [CO(2)] may be similar in FACE and enclosure studies, but there are major quantitative differences: trees were more responsive than other functional types; C(4) species showed little response; and the reduction in plant nitrogen was small and largely accounted for by decreased Rubisco. The results from this review may provide the most plausible estimates of how plants in their native environments and field-grown crops will respond to rising atmospheric [CO(2)]; but even with FACE there are limitations, which are also discussed.     

Genotypic variation in the photosynthetic competence of Sorghum bicolor seedlings subjected to polyethylene glycol-mediated drought stress

Eleven varieties of Sorghum bicolor, subjected to PEG-mediated drought stress were compared for their photosynthetic performance. The varieties differed in their relative water content over a range of PEG concentrations (0-25%). CO2 assimilation, stomatal conductance and the quantum yield of PSII electron transport decreased with increasing PEG concentrations in all varieties. However the intercellular CO2 concentration showed a nonlinear PEG concentration-dependent change. At lower PEG concentrations there was a decrease in the levels of intercellular CO2 concentration in all varieties that could be attributed to stomatal closure. At higher PEG concentrations, some varieties showed an increase in the intercellular CO2 concentration, indicating an inhibition of photosynthetic activity due to non-stomatal effects, while others did not. It was seen that the varieties differed in the stress thresholds at which stomatal and metabolic limitations to photosynthesis occur. These differences in the photosynthetic adaptation of Sorghum varieties could be useful in identifying genotypes showing large differences in photosynthetic adaptation, which could be useful in mapping photosynthetic traits for drought stress tolerance.     

Interactive Effects of Elevated CO2 and Growth Temperature on the Tolerance of Photosynthesis to Acute Heat Stress in C3 and C4 Species

Determining effects of elevated CO2 on the tolerance of photosynthesis to acute heat-stress (heat wave) is necessary for predicting plant responses to global warming, as photosynthesis is thermolabile and acute heat-stress and atmospheric CO2 will increase in the future. Few studies have examined this, and past results are variable, which may be due to methodological variation. To address this, we grew two C3 and two C4 species at current or elevated CO2 and three different growth temperatures (GT). We assessed photosynthetic thermotolerance in both unacclimated (basal tolerance) and preheat-stressed (preHS = acclimated) plants. In C3 species, basal thermotolerance of net photosynthesis (Pn) was increased In high CO2, but in C4 species, Pn thermotlerance was decreased by high CO2 (except Zea maya at low GT); CO2 effects in preHS plants were mostly small or absent, though high CO2 was detrimental in one C3 and one C4 species at warmer GT. Though high CO2 generally decreased stomatal conductance, decreases in Pn during heat stress were mostly due to non-stomatal effects. Photosystem II (PSII) efficiency was often decreased by high CO2 during heat stress, especially at high GT; CO2 effects on post-PSll electron transport were variable. Thus, high CO2 often affected photosynthetic theromotolerance, and the effects varied with photosynthetic pathway, growth temperature, and acclimation state. Most importantly, in heat-stressed plants at normal or warmer growth temperatures, high CO2 may often decrease, or not benefit as expected, tolerance of photosynthesis to acute heat stress. Therefore, interactive effects of elevated CO2 and warmer growth temperatures on acute heat tolerance may contribute to future changes in plant productivity, distribution, and diversity.     

北方粳稻光合速率、气孔导度对光强和CO2浓度的响应

 以东北地区主栽的粳稻（Oryza sativa var. japonica）品种为对象,用美国LI-cor公司生产的Li 6400光合作用测定仪控制光强、CO2浓度和温度等环境条件,阐述了光合作用和气孔导度对光和CO2浓度的响应特征及其耦合关系。结果表明,光合速率随光强或CO2浓度的提高而增大,均遵循米氏响应;在不同CO2浓度下,表观量子效率随CO2浓度的提高而增大,但CO2浓度达到800 μmol•mol－1以上时,表观量子效率有所减小;在不同光强下,表观羧化效率也随光的增强而增大,但光强达到1 600 μmol•m-2•s-1以上时,表观羧化效率也有所减小;在光强和CO2浓度协同作用下,光合速率的响应遵循双底物的米氏方程,在光强和CO2浓度均趋于饱和时,北方粳稻（品种：辽粳294）剑叶的潜在最大光合速率为71.737 8 μmol•m-2•s-1,表观量子效率为0.056 0 μmolCO2•μmol-1 photons,表观羧化效率为0.103 1 μmol•m-2•s-1/μmol•mol－1。气孔导度也随光的增强而增大,对光强的响应规律也可以用Michaelis-Menten曲线模拟,而叶面CO2浓度的提高会使气孔导度减小,气孔导度（Gs）对叶面CO2浓度（Cs）的响应可以用Gs=Gmax,c/(1+Cs/Cs0)的双曲线方程模拟。在光强(PFD)和CO2浓度协同作用下,气孔导度可以用式Gs=Gmax(PFD/PFDc)/［(1+PFD/PFDc)(1+Cs/Cs0)］+Gct估算,当CO2浓度趋于0而光强趋于饱和时,北方粳稻的潜在最大气孔导度（Gmax）为0.670 9 mol•m-2•s-1。在光强和CO2浓度协同作用下,Ball-Berry模型及其修正形式依然能很好地表达气孔导度-光合速率的耦合关系,并且用叶面饱和水汽压差（Ds）修正耦合关系中的相对湿度可以提高模拟精度。     

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.