A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Changes in stomatal conductance and net photosynthesis during phenological development in spring wheat: implications for gas exchange modelling

Gas exchange was measured from 1 month before the onset of anthesis until the end of grain filling in field-grown spring wheat, Triticum aestivum L., cv. Vinjett, in southern Sweden. Two g _s models were parameterised using these data: one Jarvis-type multiplicative g _s model (J-model), and one combined stomatal-photosynthesis model (L-model). In addition, the multiplicative g _s model parameterisation for wheat used within the European Monitoring and Evaluation Programme (EMEP-model) was tested and evaluated. The J-model performed well (R ²=0.77), with no systematic pattern of the residuals plotted against the driving variables. The L-model explained a larger proportion of the variation in g _s data when observations of A _n were used as input data (R ²=0.71) compared to when A _n was modelled (R ²=0.53). In both cases there was a systematic model failure, with g _s being over- and underestimated before and after anthesis, respectively. This pattern was caused by the non-parallel changes in g _s and A _n during plant phenological development, with A _n both peaking and starting to decline earlier as compared to g _s . The EMEP-model accounted for 41% of the variation in g _s data, with g _s being underestimated after anthesis. We conclude that, under the climatic conditions prevailing in southern Scandinavia, the performance of the combined stomatal-photosynthesis approach is hampered by the non-parallel changes in g _s and A _n, and that the phenology function of the EMEP-model, having a sharp local maximum at anthesis, should be replaced by a function with a broad non-limiting period after anthesis.     

A coupled model of stomatal conductance and photosynthesis for winter wheat

The model couples stomatal conductance (g _s) and net photosynthetic rate (P _N) describing not only part of the curve up to and including saturation irradiance (I _max), but also the range above the saturation irradiance. Maximum stomatal conductance (g _smax) and I _max can be calculated by the coupled model. For winter wheat (Triticum aestivum) the fitted results showed that maximum P _N (P _max) at 600 μmol mol⁻¹ was more than at 350 μmol mol⁻¹ under the same leaf temperature, which can not be explained by the stomatal closure at high CO₂ concentration because g _smax at 600 μmol mol⁻¹ was less than at 350 μmol mol⁻¹. The irradiance-response curves for winter wheat had similar tendency, e.g. at 25 °C and 350 μmol mol⁻¹ both P _N and g _s almost synchronously reached the maximum values at about 1 600 μmol m⁻² s⁻¹. At 25 °C and 600 μmol mol⁻¹ the I _max corresponding to P _max and g _smax was 2 080 and 1 575 μmol m⁻² s⁻¹, respectively.     

Suitability of a combined stomatal conductance and photosynthesis model for calculation of leaf-level ozone fluxes

Currently, the most important source of uncertainty in stomatal ozone flux ( FO3) modelling is the stomatal conductance ( gst) factor. Hence FO3 model accuracy will strongly depend on the gst model being implemented. In this study the recently developed semi-empirical Gst model of Dewar was coupled to the widely known biochemical photosynthesis ( An) model of Farquhar. The Gst performance of this model combination was evaluated with a 4-month time series of beech ( Fagus sylvatica L.) measurements. The Gst model was hereto optimized in two steps to a 4-day and a 8-day period. A comparison between the modelled and measured gst to O(3) (gstO3) revealed a rather good overall performance (R(2)=0.77). Errors between the model combination and the measurements are thought to be largely caused by a moderate performance of the AN model, due to poor parameterization. Two 2-day periods with distinctly differing soil and meteorological conditions were chosen to give a picture of the daily gst performance. Although instant relative differences between modelled and measured gstO3 are sometimes high, the model combination is able to simulate the rough daily courses of gstO3 and hence FO3 reasonably well. Further improvement on full parameterization of the gst model and a well-parameterized An model to be linked to are needed to draw founded conclusions about its performance. Future efforts hereto are certainly justified since the model's mechanistic nature makes it a tool able to model gst variation in space and time, O(3) effects on gst, and effective FO3.     

A high-performance system of multiple gas-exchange chambers with a laser spectrometer to estimate leaf photosynthesis,stomatal conductance,and mesophyll conductance

Journal of Plant Research - Direct measurements of ecophysiological processes such as leaf photosynthesis are often hampered due to the excessive time required for gas-exchange measurements and the...     

Variability of photosynthetic gas exchange parameters, dark respiration, and stomatal numbers in species of Polygonum

Within the genus Polygonum a large variation was found between species with regard to stomatal number, gas phase resistance, intracellular resistance and dark respiration. Interspecific variation in CO₂ compensation concentration and intercellular CO₂ concentration at constant external concentration were comparatively small. Correlations were found between stomatal number and gas phase resistance, stomatal number and Γ, and Γ and the product of dark respiration rate and intracellular resistance. The influence of dark respiration and stomatal number on photosynthetic gas exchange is discussed. It was concluded that dark respiration in light was enhanced by 22% as a mean value in 9 Polygonum species and by 62% in Polygonum lapathifolium .     

Ozone exposure causes a decoupling of conductance and photosynthesis: implications for the Ball-Berry stomatal conductance model

Industrialization has significantly altered atmospheric chemistry by increasing concentrations of chemicals such as nitrogen oxides (NO( x )) and volatile organic carbon, which react in the presence of sunlight to produce tropospheric ozone (O(3)). Ozone is a powerful oxidant that causes both visual and physiological damage to plants, impairing the ability of the plant to control processes like photosynthesis and transpiration. Damage to photosynthesis and stomatal conductance does not always occur at the same rate, which generates a problem when using the Ball-Berry model to predict stomatal conductance because the calculations directly rely on photosynthesis rates. The goals of this work were to develop a modeling framework to modify Ball-Berry stomatal conductance predictions independently of photosynthesis and to test the framework using experimental data. After exposure to elevated O(3) in open-top chambers, photosynthesis and stomatal conductance in tulip poplar changed at different rates through time. We were able to accurately model observed photosynthetic and stomatal conductance responses to chronic O(3) exposure in a Ball-Berry framework by adjusting stomatal conductance in addition to photosynthesis. This led to a significant improvement in the modeled ability to predict both photosynthesis and stomatal conductance responses to O(3).     

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.     

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.     

Exogenously-sourced ethylene increases stomatal conductance, photosynthesis, and growth under optimal and deficient nitrogen fertilization in mustard

In order to ascertain the stomatal and photosynthetic responses of mustard to ethylene under varying N availability, photosynthetic characteristics of mustard grown with optimal (80 mg N kg(-1) soil) or low (40 mg N kg(-1) soil) N were studied after the application of an ethylene-releasing compound, ethephon (2-chloroethyl phosphonic acid) at 40 days after sowing (DAS). The availability of N influenced ethylene evolution and affected stomatal conductance and photosynthesis. The effect of ethylene was smaller under deficient N where plants contained higher glucose (Glc) sensitivity, despite high ethylene evolution even in the absence of ethephon, potentially because the plants were less sensitive to ethylene per se. Ethephon application at each level of N increased ethylene and decreased Glc sensitivity, which increased photosynthesis via its effect on the photosynthetic machinery and effects on stomatal conductance. Plants grown with sufficient-N and treated with 200 μl l(-1) ethephon exhibited optimal ethylene, the greatest stomatal conductance and photosynthesis, and growth. These plants made maximum use of available N and exhibited the highest nitrogen-use efficiency (NUE).     

The effects of soil and atmospheric drought on photosynthesis and stomatal control of gas exchange in three coniferous species

The responses of steady-state CO₂ assimilation rate (A), transpiration rate (E), and stomatal conductance (g_s) to changes in leaf-to-air vapour pressure difference (δW) on one hand and to increasing soil drought on the other hand were examined in 2-year-old seedlings of Pseudotsuga menziesii, Pseudotsuga macrocarpa and Cedrus atlantica. Analysing the data through A vs intercellular CO₂ molar fraction (c_i) graphs, we could determine stomatal and mesophyll contributions to changes in A as δW or soil drought were increased. Increasing soil drought affected g_s and mesophyll photosynthesis independently, since clearly distinct predawn leaf water potential (ψ_p) regions appeared in which either stomatal or mesophyll effects prevailed for explaining the changes in A. The two Pseudotsuga species exhibited a large ψ_P range (between ca -0.8 and -1.5 to -1.9 MPa) in which only stomata were responsible for the decrease in A. A dramatic decline in mesophyll photosynthesis was noticed starting from values as high as -1.2 MPa ( C. atlantica ), -1.5 MPa ( P. macrocarpa ) and -1.9 MPa ( P. menziesii ). Increasing ΔW at high soil water content led to a sharp decline in A primarily due to an alteration of mesophyll photosynthesis. Stomatal conductance for CO₂ diffusion was affected in a lesser extent and in close correlation with the changes in mesophyll photosynthesis, which could suggest the existence of a functional linkage between mesophyll photosynthesis and stomata. Surprisingly, the drought resistant P. macrocarpa exhibited the least conservative water use efficiency in response to the two types of drought. In this species drought adaptation seems to be mainly due to its high root growth and soil prospection ability.     

NaCl胁迫对两种番茄气孔特征、气体交换参数和生物量的影响

为探讨番茄叶片气孔特征、气体交换参数和生物量对盐胁迫的响应机理,以赛棚和阿拉姆番茄为试材,通过向水培营养液中添加NaCl(0.1 mol·L^-1),在人工气候箱条件下进行为期90 d的NaCl盐胁迫处理.结果表明: NaCl胁迫导致赛棚番茄叶片的气孔密度、气孔宽度、气孔面积和气孔面积指数显著降低,降幅分别为32%、45%、25%、49%,但没有改变阿拉姆番茄叶片的气孔特征参数.同时,NaCl胁迫还导致赛棚和阿拉姆叶片气孔规则分布的空间尺度分别减少30%和43%,且赛棚品种的单个气孔最小邻域距离在盐胁迫下增加20%.另外,赛棚和阿拉姆叶片的净光合速率(P_n)、气孔导度(g_s)和蒸腾速率(T_r)在盐胁迫下均显著下降,通过气孔限制值分析发现,盐胁迫导致赛棚番茄叶片光合速率下降主要是由气孔限制因素引起的,而阿拉姆叶片则以非气孔限制因素为主导作用.盐胁迫还导致赛棚和阿拉姆番茄生物量显著降低,且地下生物量的下降幅度大于地上生物量.综合分析表明,阿拉姆的抗盐能力高于赛棚.     

Desert dogma revisited: coupling of stomatal conductance and photosynthesis in the desert shrub, Larrea tridentata

The success of the desert shrub Larrea tridentata (creosotebush) has been largely attributed to temperature acclimation and stomatal control of photosynthesis (A) under drought stress. However, there is a paucity of field data on these relationships. To address this void, we conducted a joint field and modelling study that encompassed a diverse set of environmental conditions. At a Larrea‐dominated site in southern New Mexico we manipulated soil moisture during the growing season over a 2‐year period and measured plant pre‐dawn water potential (Ψ_pd), stomatal conductance (g) and A of individual shrubs. We used these data to develop a semi‐mechanistic photosynthesis model (A–Season) that explicitly couples internal CO₂ (C_i) and g. Vapour pressure deficit (VPD) and Ψ_pd affect instantaneous g in a manner that is consistent with a biophysical model of stomatal regulation of leaf water potential. C_i is modelled as a function of g, derived from a simplification of a typical A–C_i curve. After incorporating the effects of growing temperature on stomatal behaviour, the model was able to capture the large diurnal fluctuations in A, g and C_i and the observed hysteresis in g versus C_i dynamics. Our field data and application of the A–Season model suggest that dogma attributed to Larrea's success is supported with regard to stomatal responses to VPD and Ψ_pd, but not for mechanisms of temperature acclimation and CO₂ demand.     

A fast DNA sequence handling program for Apple II computer in BASIC and 6502 assembler

A fast general purpose DNA handling program has been developedin BASIC and machine language. The program runs on the AppleII plus or on the Apple IIe microcomputer, without additionalhardware except for disk drives and printer. The program allowsfile insertion and editing, translation into protein sequence,reverse translation, search for small strings and restrictionenzyme sites. The homology may be shown either as a comparisonof two sequences or through a matrix on screen. Two additionalfeatures are: (i) drawing restriction site maps on the printer;and (ii) simulating a gel electrophoresis of restriction fragmentsboth on screen and on paper. All the operations are very fast.The more common tasks are carried out almost instantly; onlymore complex routines, like finding homology between large sequencesor searching and sorting all the restriction sites in a longsequence require longer, but still quite acceptable, times (generallyunder 30 s). Received on October 25, 1984; accepted on December 7, 1984     

The effect of exogenous abscisic acid on stomatal development, stomatal mechanics, and leaf gas exchange in Tradescantia virginiana

Gas exchange parameters and stomatal physical properties were measured in Tradescantia virginiana plants grown under well-watered conditions and treated daily with either distilled water (control) or 3.0 mM abscisic acid (ABA). Photosynthetic capacity (CO(2) assimilation rate for any given leaf intercellular CO(2) concentration [c(i)]) and relative stomatal sensitivity to leaf-to-air vapor-pressure difference were unaffected by the ABA treatment. However, at an ambient CO(2) concentration (c(a)) of 350 micromol mol(-1), ABA-treated plants operated with significantly lower c(i). ABA-treated plants had significantly smaller stomata and higher stomatal density in their lower epidermis. Stomatal aperture versus guard cell pressure (P(g)) characteristics measured with a cell pressure probe showed that although the form of the relationship was similar in control and ABA-treated plants, stomata of ABA-treated plants exhibited more complete closure at P(g) = 0 MPa and less than half the aperture of stomata in control plants at any given P(g). Scaling from stomatal aperture versus P(g) to stomatal conductance versus P(g) showed that plants grown under ABA treatment would have had significantly lower maximum stomatal conductance and would have operated with lower stomatal conductance for any given guard cell turgor. This is consistent with the observation of lower c(i)/c(a) in ABA-treated plants with a c(a) of 350 micromol mol(-1). It is proposed that the ABA-induced changes in stomatal mechanics and stomatal conductance versus P(g) characteristics constitute an improvement in water-use efficiency that may be invoked under prolonged drought conditions.     

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.     

Modelling photosynthesis in fluctuating light with inclusion of stomatal conductance, biochemical activation and pools of key photosynthetic intermediates

Photosynthetic carbon gain in rapidly fluctuating light is controlled by stomatal conductance, activation of ribulose-1,5-bisphosphate carboxylase-oxygenase, a fast induction step in the regeneration of ribulose-1,5-bisphosphate, and the build-up of pools of photosynthetic intermediates that allow post-illumination CO₂ fixation. Experimental work over recent years has identified and characterised these factors. A physiologically-based dynamic model is described here that incorporates these factors and allows the simulation of carbon gain in response to any arbitrary sequence of light levels. The model output is found to conform well to previously reported plant responses of Alocasia macrorrhiza (L.) G. Don. observed under widely differing conditions. The model shows (i) responses of net assimilation rate and stomatal conductance to constant light levels and different CO₂ concentrations that are consistent with experimental observations and predictions of a steady-state model; (ii) carbon gain to continue after the end of lightflecks, especially in uninduced leaves; (iii) carbon gain to be only marginally reduced during low-light periods of up to 2 s; (iv) a fast-inducing component in the regeneration of ribulose-1,5-bisphosphate to be limiting for up to 60 s after an increase in light in uninduced leaves: the duration of this limitation lengthens with increasing CO₂ concentration and is absent at low CO₂ concentration; (v) oxygen evolution to exceed CO₂ fixation during the first few seconds of a lightfleck, but CO₂ fixation to continue after the end of the lightfleck whereas oxygen evolution decreases to low-light rates immediately. The model is thus able to reproduce published responses of leaves to a variety of perturbations. This provides good evidence that the present formulation of the model includes the essential rate-determining factors of photosynthesis under fluctuating light conditions. Received: 27 January 1997 / Accepted: 15 April 1997     

A fortran IV computer program for automatic calculation of amino acid composition

A computer program is described which enables amino acid compositions to be calculated automatically and expressed in a variety of units. Calculations have been performed on protein hydrolyzates and cell extracts. The system is relatively simple and economical and could be applied to other column chromatographic analyses.     

Effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones

We examined the interactions of water and nitrogen availability by subjecting two Populus clones. Tristis and Eugenei, to five soil moisture and three soil nitrogen levels. Nitrogen application significantly increased net photosynthesis and stomatal conductance of flooded Eugenei and Tristis. The onset of flooding caused partial stomatal closure. Net photosynthesis significantly declined after a longer flooding period. Emergence of adventitious roots on the submerged portions of stems in both clones seemingly helped net photosynthesis fully recover in Eugenei and partially recover in Tristis. Under the progressive drought conditions, stomatal conductance was more sensitive to drought than net photosynthesis in both clones. Addition of nitrogen to progressively drying soil induced more stomatal closure in both clones. The highest water-use efficiency was found on the high-N/severe drought zone for Eugenei, whereas it was found on the high-N/mild to moderate drought zone for Tristis.