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.     

气孔导度模型在北方农牧交错带的适用性评价——以马铃薯和油葵为例

基于北方农牧交错带主要作物马铃薯和油葵的叶片气孔导度、净光合速率和气象因子的平行观测数据,对常用气孔导度模型(Jarvis模型、Ball-Berry模型、Leuning模型和Medlyn模型)进行了适用性评价.结果表明:马铃薯的气孔导度与净光合速率呈现较强的线性关系,而油葵气孔导度与净光合速率的线性关系较弱.对于马铃薯气孔导度,Ball-Berry模型模拟效果最佳,Leuning模型和Medlyn模型次之,Jarvis模型模拟效果最差;各模型的模拟值与观测值的均方根误差(RMSE)分别为0.0331、0.0371、0.0456和0.0794 mol·m^-2·s^-1,归一化均方根误差(NRMSE)分别为26.8%、30.0%、36.9%和64.3%,拟合度(R²)分别为0.96、0.61、0.91和0.88.对于油葵的气孔导度,Jarvis模型模拟效果略好于Ball-Berry模型、Medlyn模型和Leuning模型,其RMSE分别为0.2221、0.2534、0.2547和0.2758 mol·m^-2·s^-1,NRMSE分别为40.3%、46.0%、46.2%和50.1%,R²分别为0.38、0.22、0.23和0.20.气象因子对气孔导度作用的通径分析表明,北方农牧交错带马铃薯和油葵气孔导度日变化主要受饱和水汽压差影响.模型评价结果表明用于油葵的气孔导度模型需要进一步改进.     

The importance of mesophyll conductance in regulating forest ecosystem productivity during drought periods

Water availability is the most limiting factor to global plant productivity, yet photosynthetic responses to seasonal drought cycles are poorly understood, with conflicting reports on which limiting process is the most important during drought. We address the problem using a model‐data synthesis approach to look at canopy level fluxes, integrating twenty years of half hour data gathered by the FLUXNET network across six Mediterranean sites. The measured canopy level, water and carbon fluxes were used, together with an inverse canopy ecophysiological model, to estimate the bulk canopy conductance, bulk mesophyll conductance, and the canopy scale carbon pools in both the intercellular spaces and at the site of carboxylation in the chloroplasts. Thus the roles of stomatal and mesophyll conductance in the regulation of internal carbon pools and photosynthesis could be separated. A quantitative limitation analysis allowed for the relative seasonal responses of stomatal, mesophyll, and biochemical limitations to be gauged. The concentration of carbon in the chloroplast was shown to be a potentially more reliable estimator of assimilation rates than the intercellular carbon concentration. Both stomatal conductance limitations and mesophyll conductance limitations were observed to regulate the response of photosynthesis to water stress in each of the six species studied. The results suggest that mesophyll conductance could bridge the gap between conflicting reports on plant responses to soil water stress, and that the inclusion of mesophyll conductance in biosphere–atmosphere transfer models may improve their performance, in particular their ability to accurately capture the response of terrestrial vegetation productivity to drought.     

A coupled model of photosynthesis,stomatal conductance and transpiration for a rose leaf (Rosa hybrida L.)

The following three models were combined to predict simultaneously photosynthesis, stomatal conductance, transpiration and leaf temperature of a rose leaf: the biochemical model of photosynthesis of Farquhar, von Caemmerer and Berry (1980, Planta 149: 78-90), the stomatal conductance model of Ball, Woodrow and Berry (In: Biggens J, ed. Progress in photosynthesis research. The Netherlands: Martinus Nijhoff Publishers), and an energy balance model. The photosynthetic parameters: maximum carboxylation rate, potential rate of electron transport and rate of triose phosphate utilization, and their temperature dependence were determined using gas exchange data of fully expanded, young, sunlit leaves. The stomatal conductance model was calibrated independently. Prediction of net photosynthesis by the coupled model agreed well with the validation data, but the model tended to underestimate rates of stomatal conductance and transpiration. The coupled model developed in this study can be used to assist growers making environmental control decisions in glasshouse production.     

气孔导度对CO2浓度变化的模拟及其生理机制

基于气孔运动的生理生化机制重点进行了气孔导度(gs)对CO2浓度变化的响应机制分析,并推导得到气孔导度(gs)对CO2浓度变化响应模型,并以9种植物进行了模型验证。结果表明:随着CO2浓度的升高,气孔导度会逐渐降低,且下降的幅度会随着CO2浓度的升高而逐渐减弱。气孔导度对CO2浓度(Cs)变化的响应模型可以表达为gs=gmax/(1+Cs/Cs0),其中式中gmax是最大气孔导度和Cs0是实验常数。该模型较好地模拟了气孔导度随CO2浓度变化的规律,模型参数具有明确的生理意义,与Jarvis模型和Ball-Berry模型相比,该模型如何实现多种环境因子的耦合有待进一步突破。另外,模型是在短期改变叶片CO2浓度的条件下得出的,在CO2浓度长期胁迫下的适用性也有待进一步确认。     

Plantecophys - An R Package for Analysing and Modelling Leaf Gas Exchange Data

Here I present the R package ''plantecophys'', a toolkit to analyse and model leaf gas exchange data. Measurements of leaf photosynthesis and transpiration are routinely collected with portable gas exchange instruments, and analysed with a few key models. These models include the Farquhar-von Caemmerer-Berry (FvCB) model of leaf photosynthesis, the Ball-Berry models of stomatal conductance, and the coupled leaf gas exchange model which combines the supply and demand functions for CO₂ in the leaf. The ''plantecophys'' R package includes functions for fitting these models to measurements, as well as simulating from the fitted models to aid in interpreting experimental data. Here I describe the functionality and implementation of the new package, and give some examples of its use. I briefly describe functions for fitting the FvCB model of photosynthesis to measurements of photosynthesis-CO₂ response curves (''A-C_i curves''), fitting Ball-Berry type models, modelling C3 photosynthesis with the coupled photosynthesis-stomatal conductance model, modelling C4 photosynthesis, numerical solution of optimal stomatal behaviour, and energy balance calculations using the Penman-Monteith equation. This open-source package makes technically challenging calculations easily accessible for many users and is freely available on CRAN.     

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].     

A hydromechanical and biochemical model of stomatal conductance

A mathematical model of stomatal conductance is presented. It is based on whole‐plant and epidermal hydromechanics, and on two hypotheses: (1) the osmotic gradient across guard cell membranes is proportional to the concentration of ATP in the guard cells; and (2) the osmotic gradient that can be sustained per unit of ATP is proportional to the turgor pressure of adjacent epidermal cells. In the present study, guard cell [ATP] is calculated using a previously published model that is based on a widely used biochemical model of C₃ mesophyll photosynthesis. The conductance model for Vicia faba L. is parameterized and tested As with most other stomatal models, the present model correctly predicts the stomatal responses to variations in transpiration rate, irradiance and intercellular CO₂. Unlike most other models, however, this model can predict the transient stomatal opening often observed before conductance declines in response to decreases in humidity, soil water potential, or xylem conductance. The model also explicitly accommodates the mechanical advantage of the epidermis and correctly predicts that stomata are relatively insensitive to the ambient partial pressure of oxygen, as a result of the assumed dependence on ATP concentration.     

Stomatal conductance and transpirational responses of field-grown cotton to ozone

Abstract Stomatal conductance and transpiration were measured on normally-irrigated (NI) and water-stressed (WS) field-grown cotton (Gossypium hirsutum L.) exposed throughout the growing season to a gradient of ozone (O₃) concentrations. Environmental conditions during the growing season strongly affected stomatal responses and yield reductions due to O₃ exposure. Maximum stomatal conductance and transpiration decreased with increased O₃ concentration both in NI and WS treatments. Maximum conductance in severely O₃-stressed plants averaged 30% lower than charcoal-filtered (control) plants, but maximum transpiration was only 17% lower. Conductance in WS plots averaged 22% lower than in NI plots but transpiration rates were the same in both treatments. Yield reductions induced by O₃ were highly correlated (r²= 0.84) with daily transpiration. Stomata of O₃-stressed plants opened and closed at the same rate as control plants in response to changes in light intensity, suggesting that the mechanism of stomatal movement had not been impaired by exposure to O₃. Reductions in conductance and transpiration in O₃-stressed plants were attributed to inhibition of photosynthesis by O₃, leading to accumulation of CO₂ in intercellular spaces.     

The contribution of photosynthesis to the red light response of stomatal conductance

To determine the contribution of photosynthesis on stomatal conductance, we contrasted the stomatal red light response of wild-type tobacco (Nicotiana tabacum 'W38') with that of plants impaired in photosynthesis by antisense reductions in the content of either cytochrome b(6)f complex (anti-b/f plants) or Rubisco (anti-SSU plants). Both transgenic genotypes showed a lowered content of the antisense target proteins in guard cells as well as in the mesophyll. In the anti-b/f plants, CO(2) assimilation rates were proportional to leaf cytochrome b(6)f content, but there was little effect on stomatal conductance and the rate of stomatal opening. To compare the relationship between photosynthesis and stomatal conductance, wild-type plants and anti-SSU plants were grown at 30 and 300 micromol photon m(-2) s(-1) irradiance (low light and medium light [ML], respectively). Growth in ML increased CO(2) assimilation rates and stomatal conductance in both genotypes. Despite the significantly lower CO(2) assimilation rate in the anti-SSU plants, the differences in stomatal conductance between the genotypes were nonsignificant at either growth irradiance. Irrespective of plant genotype, stomatal density in the two leaf surfaces was 2-fold higher in ML-grown plants than in low-light-grown plants and conductance normalized to stomatal density was unaffected by growth irradiance. We conclude that the red light response of stomatal conductance is independent of the concurrent photosynthetic rate of the guard cells or of that of the underlying mesophyll. Furthermore, we suggest that the correlation of photosynthetic capacity and stomatal conductance observed under different light environments is caused by signals largely independent of photosynthesis.     

Phototropins but not cryptochromes mediate the blue light-specific promotion of stomatal conductance, while both enhance photosynthesis and transpiration under full sunlight

Leaf epidermal peels of Arabidopsis (Arabidopsis thaliana) mutants lacking either phototropins 1 and 2 (phot1 and phot2) or cryptochromes 1 and 2 (cry1 and cry2) exposed to a background of red light show severely impaired stomatal opening responses to blue light. Since phot and cry are UV-A/blue light photoreceptors, they may be involved in the perception of the blue light-specific signal that induces the aperture of the stomatal pores. In leaf epidermal peels, the blue light-specific effect saturates at low irradiances; therefore, it is considered to operate mainly under the low irradiance of dawn, dusk, or deep canopies. Conversely, we show that both phot1 phot2 and cry1 cry2 have reduced stomatal conductance, transpiration, and photosynthesis, particularly under the high irradiance of full sunlight at midday. These mutants show compromised responses of stomatal conductance to irradiance. However, the effects of phot and cry on photosynthesis were largely nonstomatic. While the stomatal conductance phenotype of phot1 phot2 was blue light specific, cry1 cry2 showed reduced stomatal conductance not only in response to blue light, but also in response to red light. The levels of abscisic acid were elevated in cry1 cry2. We conclude that considering their effects at high irradiances cry and phot are critical for the control of transpiration and photosynthesis rates in the field. The effects of cry on stomatal conductance are largely indirect and involve the control of abscisic acid levels.     

植物气孔导度的机理模型

Ball-Berry气孔导度模型及其修正模型是评价植物叶片气孔调节的重要工具。该文从CO₂分子在叶片气孔中扩散这个最基本的物理过程出发, 应用物理学中的分子扩散和碰撞理论、流体力学与植物生理学等知识, 严格推导出叶片气孔导度的机理模型。利用美国Li-Cor公司生产的Li-6400光合作用测定仪控制CO₂浓度、湿度和温度, 测量了华北平原冬小麦(Triticum aestivum)的光响应数据和气孔导度数据。拟合结果表明: 推导的气孔导度机理模型较之Ball-Berry气孔导度模型和Tuzet等气孔导度模型, 能更好地描述冬小麦的气孔导度与净光合速率之间的关系。如果用气孔导度的机理模型耦合光合作用对光响应的修正模型, 则耦合模型可以很好地描述华北平原冬小麦叶片气孔导度对光强的响应曲线, 并可直接估算冬小麦的最大气孔导度和对应的饱和光强, 同时可以研究最大气孔导度是否与最大净光合速率同步的问题。拟合结果还表明: 冬小麦在30 ℃、560 μmol·mol^-1CO₂, 或在32 ℃、370 μmol·mol^-1CO₂条件下, 最大气孔导度与最大净光合速率并不同步。     

Experimental Study and Mathematical Simulation of Water Use Efficiency in Wheat Leaves Affected by Some Environmental Factors

By incorporating Ball-Berry model of stomatal conductance into the models of photosynthesis and transpiration, a model of leaf water use efficiency (WUE) as affected by several environmental variables [irradiance (Ⅰ), vapor pressure deficit (VPD) and atmospheric CO2 concentration (Ca) ] was constructed. Because the environmental variables influenced the photosynthetic rate and transpiration rate in different ways, the changes of leaf WUE with these factors were quite complicated. The rates of photosynthesis and transpiration of wheat leaves were also measured in the phytotron where the environmental variables were kept within certain ranges, and leaf WUE was calculated therefrom. The results of simulation fit quite well with the measurements except at high Ca.     

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。     

Effects of elevated ozone on photosynthesis and stomatal conductance of two soybean varieties: a case study to assess impacts of one component of predicted global climate change

Global climatic change scenarios predict a significant increase in future tropospheric ozone (O₃) concentrations. The present investigation was done to assess the effects of elevated O₃ (70 and 100 ppb) on electron transport, carbon fixation, stomatal conductance and pigment concentrations in two tropical soybean ( Glycine max L.) varieties, PK 472 and Bragg. Plants were exposed to O₃ for 4 h·day⁻¹ from 10:00 to 14:00 from germination to maturity. Photosynthesis of both varieties were adversely affected, but the reduction was higher in PK 472 than Bragg. A comparison of chlorophyll a fluorescence kinetics with carbon fixation suggested greater sensitivity of dark reactions than light reactions of photosynthesis to O₃ stress. The O₃-induced uncoupling between photosynthesis and stomatal conductance in PK 472 suggests the reduction in photosynthesis may be attributed to a factor other than reduced stomatal conductance. An increase in internal CO₂ concentration in both O₃-treated soybean varieties compared suggests that the reduction in photosynthesis was due to damage to the photosynthetic apparatus, leading to accumulation of internal CO₂ and stomatal closure. The adverse impact of O₃ stress increased at higher O₃ concentrations in both soybean varieties leading to large reductions in photosynthesis. This study suggests that O₃-induced reductions in photosynthesis in tropical and temperate varieties are similar.     

Gas exchange in paphiopedilum: lack of chloroplasts in guard cells correlates with low stomatal conductance

Net photosynthesis and stomatal conductance were measured in attached leaves of Paphiopedilum insigne. At 20°C and a vapor-pressure deficit of 0.5 kilopascal, both net photosynthesis and stomatal conductance were light-saturated below 0.2 millimole per square meter per second, a response typical of shade plants. The absolute values of photosynthetic rate and conductance however were remarkably low, presumably reflecting an adaptation to the low-light, limited-nutrient habitat characteristic of these orchids. The leaves also showed a vapor-pressure deficit response, with net photosynthesis and conductance varying over a 2-fold range between 0.3 and 1.6 kilopascals.

These results confirm that Paphiopedilum stomata are functional. The correlation between achlorophyllous guard cells and low conductance rates, however, singles them out as an exceptional biological system, exhibiting basic differences from typical stomata in higher plants. Available evidence showing that guard-cell chloroplasts are needed to sustain high conductance rates at moderate to high irradiances indicates that the genetic changes leading to the loss of chloroplast differentiation in Paphiopedilum guard cells were not deleterious because of the low conductance rates characteristic of this genus.

     

Effect of water deficits on transpiration, photosynthesis and leaf conductance in cassava

Transpiration, net photosynthesis and leaf conductance decreased when leaf water potential dropped below -0.30 MPa. Both transpiration and net photosynthesis rates were considerably reduced before the leaves were visibly wilted at -0.95 MPa. Consequently, visual symptoms are unlikely to provide a useful index for characterizing water deficits in cassava ( Manihot esculenta Crantz cv. Llanera). Decreases in net photosynthesis closely followed decreases in transpiration and this suggests that stomatal closure controls both processes.     

Effect of local irradiance on CO(2) transfer conductance of mesophyll in walnut

The acclimation responses of walnut leaf photosynthesis to the irradiance microclimate were investigated by characterizing the photosynthetic properties of the leaves sampled on young trees (Juglans nigraxregia) grown in simulated sun and shade environments, and within a mature walnut tree crown (Juglans regia) in the field. In the young trees, the CO(2) compensation point in the absence of mitochondrial respiration (Gamma*), which probes the CO(2) versus O(2) specificity of Rubisco, was not significantly different in sun and shade leaves. The maximal net assimilation rates and stomatal and mesophyll conductances to CO(2) transfer were markedly lower in shade than in sun leaves. Dark respiration rates were also lower in shade leaves. However, the percentage inhibition of respiration by light during photosynthesis was similar in both sun and shade leaves. The extent of the changes in photosynthetic capacity and mesophyll conductance between sun and shade leaves under simulated conditions was similar to that observed between sun and shade leaves collected within the mature tree crown. Moreover, mesophyll conductance was strongly correlated with maximal net assimilation and the relationships were not significantly different between the two experiments, despite marked differences in leaf anatomy. These results suggest that photosynthetic capacity is a valuable parameter for modelling within-canopies variations of mesophyll conductance due to leaf acclimation to light.     

Responses of selected birch (Betula pendula Roth) clones to ozone change over time

A long-term free air ozone fumigation experiment was conducted to study changes in physiological ozone responses during tree ontogeny and exposure time in ozone sensitive and tolerant clones of European white birch (Betula pendula Roth), originated from south and central Finland. The trees were grown in soil in natural microclimatic conditions under ambient ozone (control) and 1.4-1.7 x ambient (elevated) ozone from May 1996 to October 2001, and were measured for stem and foliage growth, net photosynthesis, stomatal conductance, stomatal density, visible injuries, foliar starch content and bud formation. After 6 years of exposure, the magnitude of ozone-induced growth reductions in the sensitive clone was 12-48% (significant difference), levels similar or greater than those reported earlier for 2- and 3-year-old saplings undergoing shorter exposures. In the tolerant clone, growth of these larger trees was reduced by 1-38% (significant difference in stem volume), although the saplings had previously been unaffected. In both clones, ozone stress led to significantly reduced leaf-level net photosynthesis but significantly increased stomatal conductance rates during the late summer, resulting in a lower carbon gain for bud formation and the onset of visible foliar injuries. Increasing ozone sensitivity with duration of exposure was explained by a change in growth form (relatively reduced foliage mass), a lower photosynthesis to stomatal conductance ratio during the late summer, and deleterious carry-over effects arising from the reduced number of over-wintering buds.     

Physiology-phenology interactions in a productive semi-arid pine forest

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D(L)) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g(s)) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g(s )sensitivity to D(L) below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis-phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.