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.     

Leaf nitrogen, photosynthesis, conductance and transpiration: scaling from leaves to canopies

A model is presented which solves simultaneously for leaf-scale stomatal conductance, CO₂ assimilation and the energy balance as a function of leaf position within canopies of well-watered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogen- and light-use efficiency), and to set bounds on the magnitudes of the parameters. For canopies low in total N, daily assimilation rates are ～10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, G_c should be calculated as G_c=Aσ(f_slg_sl+f_shg_sh), where Δ is leaf area index, f_si and f_sh are the fractions of sunlit and shaded leaves at each level, and g_si and g_sh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the ‘big-leaf’ version of the Penman-Monteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation. Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogen- and radiation-use efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2- to 3-fold variation in these efficiencies.     

黄土高原苹果园蒸腾导度大气驱动规律比较

植物蒸腾导度是表征土壤-植物-大气连续体（SPAC）中植物-大气间水汽传导过程、反映植物水分调控能力的一类重要变量,常见有冠层导度（G_c）、冠层气孔导度（G_s）与叶片气孔导度（g_s）,明确三者在反映冠层蒸腾过程时的异同或关联性对于理解植物水分利用机制具有重要意义。本研究基于对黄土高原果园苹果树生长季内树干液流（J_s）及环境因子的连续观测,计算了G_c、G_s及脱耦联系数（Ω）等变量,并与短期连续观测的叶片气孔导度（g_s）比较,分析了G_c、G_s和g_s在反映冠层蒸腾特征方面的异同及其关系。结果表明,日变化过程中G_s、g_s呈"单峰"型曲线,而G_c则呈"先增后减,午后抬升"的"双峰"型曲线。g_s与G_s存在较紧密的线性关系（R²=0.80）,但与G_c的线性关系较弱（R²=0.02）。G_c、G_s均随大气水汽压亏缺（VPD）的变化呈现确定的规律,其中,上边界函数呈递减的对数函数关系,平均值则符合先增后减的Log-Normal函数关系（R²>0.95）,拐点对应的VPD值分别为1.33和1.16 kPa。在一日内,G_s对VPD变化的响应过程与g_s对VPDL （基于叶片温度计算的水汽压亏缺）变化的响应过程总体一致,其一致性高于G_c对VPD变化的响应。整个生长季（4-10月）中果树的Ω平均值为0.12,随着Ω递减,G_c与G_s的线性相关性愈趋紧密,其斜率呈递增趋势,G_c越来越趋近于G_s。研究结果表明,在北方地区,基于树干液流的监测能较准确的推导整株并估算林分的冠层蒸腾导度。与实测g_s的变化过程比较,G_s比G_c具有更高的一致性,G_s可以作为描述苹果树水分利用过程响应大气驱动的更为恰当的变量。     

整树水力导度协同冠层气孔导度调节森林蒸腾

冠层气孔导度决定森林的蒸腾效率,它对驱动水汽移动的水汽应力的响应受树木水力结构的影响,并随水汽压亏缺上升和水力导度下降而降低,维持水势在最低阈值之上,避免出现水力灾变,调控冠层蒸腾。由于叶形和树冠结构的特点,部分脱耦联反映了湿润地区阔叶林冠层与大气的水汽交换特征,单纯以气孔导度的变化难以完整描述水分通量的调节规律,因而,需要考虑冠层气孔导度与水力导度协同控制冠层蒸腾的潜在机理。通过整合叶片气孔气体交换、树干液流、冠层微气象和其他环境因子的野外观测值,估测不同时间尺度的森林冠层气孔导度与大气的脱耦联系数和变异范围,以基于树干液流的冠层蒸腾,结合叶片/土壤水势梯度计算的水力导度,分析水力导度影响冠层气孔导度响应水汽压亏缺的敏感性,可以揭示和阐明水力导度和冠层气孔导度联合调节森林蒸腾的机理,对准确估测全球变化背景下森林对水资源利用的潜在生态效应有明显的理论意义。     

北京山区侧柏林冠层-大气蒸腾导度模拟及环境因子响应

蒸腾导度模型是衡量冠层-大气界面水汽输出的重要阻力模型,研究其特征及对环境因子的响应,为揭示森林冠层-大气界面水汽输出阻力机制提供理论依据。以首都圈森林生态系统定位观测研究站侧柏林为研究对象,采用TDP热探针法测定侧柏林树干液流密度,同步监测光合有效辐射、饱和水汽压差、气温、风速等主要环境因子,分析冠层导度和空气动力学导度的动态变化,构建冠层-大气蒸腾导度模型并模拟,明确冠层-大气蒸腾导度对各环境因子的响应关系。结果表明:蒸腾导度季节变化表现为非生长季与冠层导度趋势一致,生长季与空气动力学导度趋势一致,全年均为单峰趋势。冬季蒸腾导度与冠层导度保持较稳定差值(45 mol m^(-2 )s^-1左右),其他季节蒸腾导度与冠层导度、空气动力学导度的最大差值,均在各季节冠层导度、空气动力学导度的峰值水平。全年日均蒸腾导度冬季最大(86.92 mol m^(-2 )s^-1),其他季节较小且稳定(40—50 mol m^(-2 )s^-1之间)。在非生长季各环境因子对蒸腾导度的影响与对冠层导度的影响基本一致,温度为主要影响因子(r=-0.198),其他环境因子影响较小(r<0.1);在生长季中风速为主要影响因子(r=0.488),光合有效辐射(r=0.228)和饱和水汽压差(r=-0.299)的影响明显升高,温度的影响降低(r=0.114)。蒸腾导度模型较好的模拟了冠层-大气界面侧柏蒸腾不同季节的变化规律,阐明了各环境因子和冠层导度、空气动力学导度对蒸腾导度的影响机制,证实在生长季应重视空气动力学导度对蒸腾的影响。     

Impact of environmental factors on the decoupling coefficient and the estimation of canopy stomatal conductance for ever-green broad-leaved tree species

精确模拟冠层气孔导度(G_S)对于评估区域蒸散具有重要意义。该研究选择两种常见的人工阔叶树种尾叶桉(Eucalyptus urophylla, 外来种)和木荷(Schima superba, 本地种)作为研究对象, 利用K?stner法和修订的Penman-Monteith公式计算冠层平均气孔导度(分别定义为G_S1和G_S2)。研究还分析了环境因子对冠层脱耦联系数(Ω)的影响, 并用其来评价两种方法模拟的冠层气孔导度的合理性。结果表明, 两个树种冠层气孔导度均与气象条件耦合较好(尾叶桉: Ω = 0.10 ± 0.03, 木荷: Ω = 0.17 ± 0.03)。主成分分析显示, 光合有效辐射(PAR)以及水汽压亏缺(D)显著影响Ω的大小, 而风速(u)的影响较小。单因素分析则发现各环境因子与Ω之间的相关性并不显著。边界线分析表明D和PAR的增加使得Ω最终趋向于一个与树种有关的稳定值(木荷≈ 0.20, 尾叶桉≈ 0.05), 而Ω随u的增加呈幂指数下降。与木荷相比, 尾叶桉具有更高的气孔导度(尾叶桉和木荷的G_S2年平均值分别为(33.42 ± 9.37) mmol·m ^-2·s ^-1和(23.40 ± 2.03) mmol·m ^-2·s ^-1), 并且尾叶桉和木荷的G_S1与G_S2的线性拟合斜率分别为0.92 (R ² ≈ 0.70)和0.98 (R ² ≈ 0.76) , 表明G_S1比G_S2高估了冠层气孔导度。另外, G_S1和G_S2对水汽压亏缺的敏感性与参比气孔导度(G_Siref, D = 1 kPa时的气孔导度)的比值Pi与Ω紧密相关。根据统计, 尾叶桉和木荷的G_S1估计值在Ω = 0.05-0.15 (83.1%的数据)和0.10-0.20 (47.8%的数据)之间时是相对可靠的。     

Estimating the sensitivity of stomatal conductance to photosynthesis: a review

A common approach for estimating fluxes of CO₂ and water in canopy models is to couple a model of photosynthesis (A_n) to a semi‐empirical model of stomatal conductance (g_s) such as the widely validated and utilized Ball–Berry (BB) model. This coupling provides an effective way of predicting transpiration at multiple scales. However, the designated value of the slope parameter (m) in the BB model impacts transpiration estimates. There is a lack of consensus regarding how m varies among species or plant functional types (PFTs) or in response to growth conditions. Literature values are highly variable, with inter‐species and intra‐species variations of >100%, and comparisons are made more difficult because of differences in collection techniques. This paper reviews the various methods used to estimate m and highlights how variations in measurement techniques or the data utilized can influence the resultant m. Additionally, this review summarizes the reported responses of m to [CO₂] and water stress, collates literature values by PFT and compiles nearly three decades of values into a useful compendium.     

Effect of black bean aphid, Aphis fabae, on transpiration, stomatal conductance and crude protein content of faba bean

This study was initiated to investigate effects of damage by 0, 5 and 10 aphids/plant on the physiology of faba bean plants throughout different feeding periods and at two plant development stages. Immediately following removal of Aphis fabae, measurements showed 84–229% increase in transpiration rate. These changes were proportional to the number of aphids and infestation duration. Injury by A. fabae caused the stomatal conductance to be much higher in the leaves of infested plants. Leaf stomatal conductance of the infested plants increased significantly by 51–224% depending on initial aphid densities and feeding intervals. This increase was proportional to the infestation level for each date. Length of infestation period and plant growth stage seemed to have no clear effect on stomatal apertures. Aphid feeding caused a damage of about 7–33% of crude protein levels in the leaf tissue. This reduction increased with increasing infestation levels and time, except for 28‐day‐old plants on 28 days. The physiological effects of aphid feeding on water vapour and chemical composition of damaged leaves are particularly serious when the population is high.     

黄土高原油松冠层气孔导度和蒸腾变化特征与模拟

油松是黄土高原重要的造林树种,模拟其冠层气孔导度和蒸腾对区域水量平衡计算和人工林可持续经营具有重要意义。基于2015—2018年TDP(Thermal dissipation probes)方法所测得液流数据,分析了黄土高原地区油松冠层平均气孔导度(g_c)与冠层蒸腾(Tr)的变化特征与影响因素,并采用Penman-Monteith公式和Jarvis型气孔导度模型模拟了其g_c和Tr的变化过程,结果表明：(1)该地区油松g_c和Tr日内变化均呈现单峰型,日均蒸腾耗水量为(1.25±0.57) mm/d,生长季(4—10月)总蒸腾耗水量均值为195.47 mm。(2)g_c的日内变化受太阳辐射(Rad)驱动(偏相关系数为0.65),当Rad高于300 W/m²时,驱动作用减弱;g_c的日内变化受水汽压亏缺(VPD)控制(偏相关系数为-0.41),随VPD的增加而降低;g_c的日际变化受土壤水分限制(偏相关系数为0.46),当根区相对有效含水率(RE...     

Carbonyl sulfide (COS) as a tracer for canopy photosynthesis, transpiration and stomatal conductance: potential and limitations

The theoretical basis for the link between the leaf exchange of carbonyl sulfide (COS), carbon dioxide (CO(2)) and water vapour (H(2)O) and the assumptions that need to be made in order to use COS as a tracer for canopy net photosynthesis, transpiration and stomatal conductance, are reviewed. The ratios of COS to CO(2) and H(2)O deposition velocities used to this end are shown to vary with the ratio of the internal to ambient CO(2) and H(2)O mole fractions and the relative limitations by boundary layer, stomatal and internal conductance for COS. It is suggested that these deposition velocity ratios exhibit considerable variability, a finding that challenges current parameterizations, which treat these as vegetation-specific constants. COS is shown to represent a better tracer for CO(2) than H(2)O. Using COS as a tracer for stomatal conductance is hampered by our present poor understanding of the leaf internal conductance to COS. Estimating canopy level CO(2) and H(2)O fluxes requires disentangling leaf COS exchange from other ecosystem sources/sinks of COS. We conclude that future priorities for COS research should be to improve the quantitative understanding of the variability in the ratios of COS to CO(2) and H(2)O deposition velocities and the controlling factors, and to develop operational methods for disentangling ecosystem COS exchange into contributions by leaves and other sources/sinks. To this end, integrated studies, which concurrently quantify the ecosystem-scale CO(2), H(2)O and COS exchange and the corresponding component fluxes, are urgently needed.     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

12.

环境因子对常绿阔叶树种脱耦联系数及冠层气孔导度估算的影响   总被引：1，自引：0，他引：1       下载免费PDF全文

张振振  赵平  赵秀华  张锦秀  朱丽薇  欧阳磊  张笑颜 《植物生态学报》2018,42(12):1179-1191

精确模拟冠层气孔导度(G_S)对于评估区域蒸散具有重要意义。该研究选择两种常见的人工阔叶树种尾叶桉(Eucalyptus urophylla, 外来种)和木荷(Schima superba, 本地种)作为研究对象, 利用K?stner法和修订的Penman-Monteith公式计算冠层平均气孔导度(分别定义为G_S1和G_S2)。研究还分析了环境因子对冠层脱耦联系数(Ω)的影响, 并用其来评价两种方法模拟的冠层气孔导度的合理性。结果表明, 两个树种冠层气孔导度均与气象条件耦合较好(尾叶桉: Ω = 0.10 ± 0.03, 木荷: Ω = 0.17 ± 0.03)。主成分分析显示, 光合有效辐射(PAR)以及水汽压亏缺(D)显著影响Ω的大小, 而风速(u)的影响较小。单因素分析则发现各环境因子与Ω之间的相关性并不显著。边界线分析表明D和PAR的增加使得Ω最终趋向于一个与树种有关的稳定值(木荷≈ 0.20, 尾叶桉≈ 0.05), 而Ω随u的增加呈幂指数下降。与木荷相比, 尾叶桉具有更高的气孔导度(尾叶桉和木荷的G_S2年平均值分别为(33.42 ± 9.37) mmol·m ^-2·s ^-1和(23.40 ± 2.03) mmol·m ^-2·s ^-1), 并且尾叶桉和木荷的G_S1与G_S2的线性拟合斜率分别为0.92 (R ² ≈ 0.70)和0.98 (R ² ≈ 0.76) , 表明G_S1比G_S2高估了冠层气孔导度。另外, G_S1和G_S2对水汽压亏缺的敏感性与参比气孔导度(G_Siref, D = 1 kPa时的气孔导度)的比值Pi与Ω紧密相关。根据统计, 尾叶桉和木荷的G_S1估计值在Ω = 0.05-0.15 (83.1%的数据)和0.10-0.20 (47.8%的数据)之间时是相对可靠的。     

13.

Mesophyll conductance in land surface models: effects on photosynthesis and transpiration     

Jürgen Knauer  Snke Zaehle  Martin G. De Kauwe  Vanessa Haverd  Markus Reichstein  Ying Sun 《The Plant journal : for cell and molecular biology》2020,101(4):858-873

The CO₂ transfer conductance within plant leaves (mesophyll conductance, g_m) is currently not considered explicitly in most land surface models (LSMs), but instead treated implicitly as an intrinsic property of the photosynthetic machinery. Here, we review approaches to overcome this model deficiency by explicitly accounting for g_m, which comprises the re‐adjustment of photosynthetic parameters and a model describing the variation of g_m in dependence of environmental conditions. An explicit representation of g_m causes changes in the response of photosynthesis to environmental factors, foremost leaf temperature, and ambient CO₂ concentration, which are most pronounced when g_m is small. These changes in leaf‐level photosynthesis translate into a stronger climate and CO₂ response of gross primary productivity (GPP) and transpiration at the global scale. The results from two independent studies show consistent latitudinal patterns of these effects with biggest differences in GPP in the boreal zone (up to ~15%). Transpiration and evapotranspiration show spatially similar, but attenuated, changes compared with GPP. These changes are indirect effects of g_m caused by the assumed strong coupling between stomatal conductance and photosynthesis in current LSMs. Key uncertainties in these simulations are the variation of g_m with light and the robustness of its temperature response across plant types and growth conditions. Future research activities focusing on the response of g_m to environmental factors and its relation to other plant traits have the potential to improve the representation of photosynthesis in LSMs and to better understand its present and future role in the Earth system.     

14.

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species          下载免费PDF全文

Dongliang Xiong  Cyril Douthe  Jaume Flexas 《Plant, cell & environment》2018,41(2):436-450

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

15.

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato     

James A. Bunce 《Photosynthesis research》1988,18(3):357-362

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

16.

北京山区油松和元宝槭冠层气孔导度特征及其环境响应          下载免费PDF全文

陈胜楠  陈左司南  张志强 《植物生态学报》2021,45(12):1329-1340

叶片气孔是植物进行水汽交换的通道, 影响着植物的蒸腾和光合作用。然而叶片气孔行为受环境条件和树种类型的影响, 不同树种冠层气孔导度对环境因子响应的差异性, 以及在生长季不同时期叶片气孔对冠层蒸腾的调节作用是否会发生改变, 仍不清楚。该研究目的是通过探究各环境因子对不同树种冠层气孔导度的相对贡献率以及叶片气孔对冠层蒸腾的调节作用, 为深入了解植物水分利用状况和山区森林经营提供参考依据。于2018年生长季以北京八达岭国家森林公园内的58年生油松(Pinus tabuliformis)和39年生元宝槭(Acer truncatum)为研究对象, 利用热扩散技术对其树干液流进行连续监测, 并同步监测环境因子。利用彭曼公式计算冠层气孔导度(G_s)。主要结果: (1)油松和元宝槭日间G_s在日、月时间尺度上存在明显差异。5-7月油松和元宝槭日动态G_s均随饱和水汽压差(VPD)和太阳辐射(GR)的增加呈上升趋势, 上升持续时间比8月和9月长; 在月尺度上, 随着VPD、GR的降低和土壤湿度(VWC)的升高, G_s从5月到9月整体上升。(2)利用增强回归树法分析得到VWC和VPD对G_s的贡献率最大, 其次是GR、气温和风速。VWC和VPD对油松G_s的贡献率分别为66.4%和17.4%, 对元宝槭G_s的贡献率分别为54.8%和21.0%。(3)油松和元宝槭的dG_s/dlnVPD值与参考冠层气孔导度之间的斜率均显著高于0.6, 气孔调节作用相对较强。综上所述, 气孔对环境因子的响应在树种以及生长季不同时期之间存在差异, 为防止水分过度散失, 两树种在不同土壤水分条件下均通过严格的气孔调节控制蒸腾量。     

17.

Reconciling the optimal and empirical approaches to modelling stomatal conductance   总被引：2，自引：0，他引：2  

BELINDA E. MEDLYN  REMKO A. DUURSMA  DEREK EAMUS  DAVID S. ELLSWORTH  I. COLIN PRENTICE  CRAIG V. M. BARTON  KRISTINE Y. CROUS  PAOLO DE ANGELIS  MICHAEL FREEMAN  LISA WINGATE 《Global Change Biology》2011,17(6):2134-2144

Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long‐standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g₁, is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g₁ is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO₂, and thus can be used to test for stomatal acclimation to elevated CO₂. The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change.     

18.

Sensitivity of infrared water vapor analyzers to oxygen concentration and errors in stomatal conductance     

Bunce JA 《Photosynthesis research》2002,71(3):273-276

Use of infrared analyzers to measure water vapor concentrations in photosynthesis systems is becoming common. It is known that sensitivity of infrared carbon dioxide and water vapor analyzers is affected by the oxygen concentration in the background gas, particularly for absolute analyzers, but the potential for large errors in estimates of stomatal conductance due to effects of oxygen concentration on the sensitivity of infrared water vapor analyzers is not widely recognized. This work tested three types of infrared water vapor analyzers for changes in sensitivity of infrared water vapor analyzers depending on the oxygen content of the background gas. It was found that changing from either 0 or 2% to 21% oxygen in nitrogen decreased the sensitivity to water vapor for all three types of infrared water vapor analyzers by about 4%. The change in sensitivity was linear with oxygen mole fraction. The resulting error in calculated stomatal conductance would depend strongly on the leaf to air vapor pressure difference and leaf temperature, and also on whether leaf temperature was directly measured or calculated from energy balance. Examples of measurements of gas exchange on soybean leaves under glasshouse conditions indicated that changing from 21% to 2% oxygen produced an artifactual apparent increase in stomatal conductance which averaged about 30%. Similar errors occurred for `conductances' of wet filter paper. Such errors could affect inferences about the carbon dioxide dependence of the sensitivity of photosynthesis to oxygen. This revised version was published online in June 2006 with corrections to the Cover Date.     

19.

Effects of ozone on stomatal conductance and photosynthesis in populations of Plantago major L.   总被引：7，自引：5，他引：2  

K. REILING  A. W. DAVISON 《The New phytologist》1995,129(4):587-594

     

20.

Simple models for stomatal conductance derived from a process model: cross-validation against sap flux data     

Buckley TN  Turnbull TL  Adams MA 《Plant, cell & environment》2012,35(9):1647-1662

Representation of stomatal physiology in models of plant-atmosphere gas exchange is minimal, and direct application of process-based models is limited by difficulty of parameter estimation. We derived simple models of stomatal conductance from a recent process-based model, and cross-validated them against measurements of sap flux (176-365 d in length) in 36 individual trees of two age classes for two Eucalyptus species across seven sites in the mountains of southeastern Australia. The derived models - which are driven by irradiance and evaporative demand and have two to four parameters that represent sums and products of biophysical parameters in the process model - reproduced a median 83-89% of observed variance in half-hourly and diurnally averaged sap flux, and performed similarly whether fitted using a random sample of all data or using 1 month of data from spring or autumn. Our simple models are an advance in predicting plant water use because their parameters are transparently related to reduced processes and properties, enabling easy accommodation of improved knowledge about how those parameters respond to environmental change and differ among species.     

环境因子对常绿阔叶树种脱耦联系数及冠层气孔导度估算的影响

精确模拟冠层气孔导度(G_S)对于评估区域蒸散具有重要意义。该研究选择两种常见的人工阔叶树种尾叶桉(Eucalyptus urophylla, 外来种)和木荷(Schima superba, 本地种)作为研究对象, 利用K?stner法和修订的Penman-Monteith公式计算冠层平均气孔导度(分别定义为G_S1和G_S2)。研究还分析了环境因子对冠层脱耦联系数(Ω)的影响, 并用其来评价两种方法模拟的冠层气孔导度的合理性。结果表明, 两个树种冠层气孔导度均与气象条件耦合较好(尾叶桉: Ω = 0.10 ± 0.03, 木荷: Ω = 0.17 ± 0.03)。主成分分析显示, 光合有效辐射(PAR)以及水汽压亏缺(D)显著影响Ω的大小, 而风速(u)的影响较小。单因素分析则发现各环境因子与Ω之间的相关性并不显著。边界线分析表明D和PAR的增加使得Ω最终趋向于一个与树种有关的稳定值(木荷≈ 0.20, 尾叶桉≈ 0.05), 而Ω随u的增加呈幂指数下降。与木荷相比, 尾叶桉具有更高的气孔导度(尾叶桉和木荷的G_S2年平均值分别为(33.42 ± 9.37) mmol·m ^-2·s ^-1和(23.40 ± 2.03) mmol·m ^-2·s ^-1), 并且尾叶桉和木荷的G_S1与G_S2的线性拟合斜率分别为0.92 (R ² ≈ 0.70)和0.98 (R ² ≈ 0.76) , 表明G_S1比G_S2高估了冠层气孔导度。另外, G_S1和G_S2对水汽压亏缺的敏感性与参比气孔导度(G_Siref, D = 1 kPa时的气孔导度)的比值Pi与Ω紧密相关。根据统计, 尾叶桉和木荷的G_S1估计值在Ω = 0.05-0.15 (83.1%的数据)和0.10-0.20 (47.8%的数据)之间时是相对可靠的。     

Mesophyll conductance in land surface models: effects on photosynthesis and transpiration

The CO₂ transfer conductance within plant leaves (mesophyll conductance, g_m) is currently not considered explicitly in most land surface models (LSMs), but instead treated implicitly as an intrinsic property of the photosynthetic machinery. Here, we review approaches to overcome this model deficiency by explicitly accounting for g_m, which comprises the re‐adjustment of photosynthetic parameters and a model describing the variation of g_m in dependence of environmental conditions. An explicit representation of g_m causes changes in the response of photosynthesis to environmental factors, foremost leaf temperature, and ambient CO₂ concentration, which are most pronounced when g_m is small. These changes in leaf‐level photosynthesis translate into a stronger climate and CO₂ response of gross primary productivity (GPP) and transpiration at the global scale. The results from two independent studies show consistent latitudinal patterns of these effects with biggest differences in GPP in the boreal zone (up to ~15%). Transpiration and evapotranspiration show spatially similar, but attenuated, changes compared with GPP. These changes are indirect effects of g_m caused by the assumed strong coupling between stomatal conductance and photosynthesis in current LSMs. Key uncertainties in these simulations are the variation of g_m with light and the robustness of its temperature response across plant types and growth conditions. Future research activities focusing on the response of g_m to environmental factors and its relation to other plant traits have the potential to improve the representation of photosynthesis in LSMs and to better understand its present and future role in the Earth system.     

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

北京山区油松和元宝槭冠层气孔导度特征及其环境响应

叶片气孔是植物进行水汽交换的通道, 影响着植物的蒸腾和光合作用。然而叶片气孔行为受环境条件和树种类型的影响, 不同树种冠层气孔导度对环境因子响应的差异性, 以及在生长季不同时期叶片气孔对冠层蒸腾的调节作用是否会发生改变, 仍不清楚。该研究目的是通过探究各环境因子对不同树种冠层气孔导度的相对贡献率以及叶片气孔对冠层蒸腾的调节作用, 为深入了解植物水分利用状况和山区森林经营提供参考依据。于2018年生长季以北京八达岭国家森林公园内的58年生油松(Pinus tabuliformis)和39年生元宝槭(Acer truncatum)为研究对象, 利用热扩散技术对其树干液流进行连续监测, 并同步监测环境因子。利用彭曼公式计算冠层气孔导度(G_s)。主要结果: (1)油松和元宝槭日间G_s在日、月时间尺度上存在明显差异。5-7月油松和元宝槭日动态G_s均随饱和水汽压差(VPD)和太阳辐射(GR)的增加呈上升趋势, 上升持续时间比8月和9月长; 在月尺度上, 随着VPD、GR的降低和土壤湿度(VWC)的升高, G_s从5月到9月整体上升。(2)利用增强回归树法分析得到VWC和VPD对G_s的贡献率最大, 其次是GR、气温和风速。VWC和VPD对油松G_s的贡献率分别为66.4%和17.4%, 对元宝槭G_s的贡献率分别为54.8%和21.0%。(3)油松和元宝槭的dG_s/dlnVPD值与参考冠层气孔导度之间的斜率均显著高于0.6, 气孔调节作用相对较强。综上所述, 气孔对环境因子的响应在树种以及生长季不同时期之间存在差异, 为防止水分过度散失, 两树种在不同土壤水分条件下均通过严格的气孔调节控制蒸腾量。     

Reconciling the optimal and empirical approaches to modelling stomatal conductance

Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long‐standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g₁, is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g₁ is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO₂, and thus can be used to test for stomatal acclimation to elevated CO₂. The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change.     

Sensitivity of infrared water vapor analyzers to oxygen concentration and errors in stomatal conductance

Use of infrared analyzers to measure water vapor concentrations in photosynthesis systems is becoming common. It is known that sensitivity of infrared carbon dioxide and water vapor analyzers is affected by the oxygen concentration in the background gas, particularly for absolute analyzers, but the potential for large errors in estimates of stomatal conductance due to effects of oxygen concentration on the sensitivity of infrared water vapor analyzers is not widely recognized. This work tested three types of infrared water vapor analyzers for changes in sensitivity of infrared water vapor analyzers depending on the oxygen content of the background gas. It was found that changing from either 0 or 2% to 21% oxygen in nitrogen decreased the sensitivity to water vapor for all three types of infrared water vapor analyzers by about 4%. The change in sensitivity was linear with oxygen mole fraction. The resulting error in calculated stomatal conductance would depend strongly on the leaf to air vapor pressure difference and leaf temperature, and also on whether leaf temperature was directly measured or calculated from energy balance. Examples of measurements of gas exchange on soybean leaves under glasshouse conditions indicated that changing from 21% to 2% oxygen produced an artifactual apparent increase in stomatal conductance which averaged about 30%. Similar errors occurred for `conductances' of wet filter paper. Such errors could affect inferences about the carbon dioxide dependence of the sensitivity of photosynthesis to oxygen. This revised version was published online in June 2006 with corrections to the Cover Date.     

Simple models for stomatal conductance derived from a process model: cross-validation against sap flux data

Representation of stomatal physiology in models of plant-atmosphere gas exchange is minimal, and direct application of process-based models is limited by difficulty of parameter estimation. We derived simple models of stomatal conductance from a recent process-based model, and cross-validated them against measurements of sap flux (176-365 d in length) in 36 individual trees of two age classes for two Eucalyptus species across seven sites in the mountains of southeastern Australia. The derived models - which are driven by irradiance and evaporative demand and have two to four parameters that represent sums and products of biophysical parameters in the process model - reproduced a median 83-89% of observed variance in half-hourly and diurnally averaged sap flux, and performed similarly whether fitted using a random sample of all data or using 1 month of data from spring or autumn. Our simple models are an advance in predicting plant water use because their parameters are transparently related to reduced processes and properties, enabling easy accommodation of improved knowledge about how those parameters respond to environmental change and differ among species.