Stomatal regulation by microclimate and tree water relations: interpreting ecophysiological field data with a hydraulic plant model

Dynamics in microclimate and physiological plant traits were studied for Pubescent oak and Scots pine in a dry inner-alpine valley in Switzerland, at a 10 min resolution for three consecutive years (2001-2003). As expected, stomata tended to close with increasing drought in air and soil. However, stomatal aperture in oak was smaller than in pine under relatively wet conditions, but larger under dry conditions. To explore underlying mechanisms, a model was applied that (i) quantifies water relations within trees from physical principles (mechanistic part) and (ii) assumes that signals from light, stomatal aperture, crown water potential, and tree water deficit in storage pools control stomata (systemic part). The stomata of pine showed a more sensitive response to increasing drought because both factors, the slowly changing tree water deficit and the rapidly changing crown water potential, closed the stomata. By contrast, the stomata of oak became less drought-sensitive as the closing signal of crown water potential was opposed by the opening signal of tree water deficit. Moreover, parameter optimization suggests that oak withdrew more water from the storage pools and reduced leaf water potentials to lower levels, without risking serious damage by cavitation. The new model thus suggests how the hydraulic water flow and storage system determines the responses in stomatal aperture and transpiration to drought at time scales ranging from hours to multiple years, and why pine and oak might differ in such responses. These differences explain why oaks are more efficient competitors during drought periods, although this was not the case in the extremely dry year 2003, which provoked massive leaf loss and, from July onwards, physiological activity almost ceased.     

A model of water flow through plants incorporating shoot/root 'message' control of stomatal conductance

Abstract. A model of water flow from the soil into the plant, and from the plant to the atmosphere is described. There are three state variables in the model: the soil, root and shoot water contents. The flow rate of water from the soil to the root is calculated by dividing the gradient in water potential by a resistance, comprising the resistance from the bulk soil to the root surface, and that from the root surface to the root interior. The resistance in the soil depends on the soil hydraulic conductivity, which in turn depends on the soil water potential. The flow rate from the root to the shoot is given by the gradient in water potential divided by a resistance, which depends on the structural dry mass of the plant. Transpiration is described by the Penman-Monteith equation. The plant water characteristics can be modified to take account of osmotic and cell wall rigidity parameters. The model incorporates the concept of shoot/root ‘messages’ of water stress, which influence stomatal conductance. The message works through the generation of a hormone as the pressure potential in the shoot (mesophyll) or root falls. This hormone induces a shift of osmoticum from the guard cells to the surrounding mesophyll cells, which causes an increase (i.e. closer to zero) in the osmotic potential in these cells. This, in turn, causes a decrease in their pressure potential, and so reduces stomatal conductance. The model is used as a framework to address some of the issues that have recently been raised concerning the role of water potential in describing water flow through plants. We conclude that, with the hormone present, there is unlikely to be a unique relationship between stomatal conductance and shoot total water potential, since stomatal conductance depends on the pressure potential in the guard cells, which may differ from that in other cells. Nevertheless, this does not imply that water potential is not an important, and indeed fundamental, component for describing water flow through plants. Other aspects of water flow through plants are also considered, such as diurnal patterns of shoot, root and soil water potential components. It is seen that these may differ from the commonly held view that, as the soil dries down, they all attain the same values during the dark period, and which, as we show, is largely unsubstantiated either theoretically or experimentally.     

不同生境下入侵种假臭草叶片的气孔特征

【背景】探讨入侵种假臭草不同生境下气孔的变化规律,揭示假臭草种群在不同生境下所采取的生长对策及适应机制,可为入侵生物的防治提供参考。【方法】采取光学显微镜系统观察桉树林、木薯地、弃耕地、公路边4种生境下假臭草叶片的气孔特征。【结果】光照和土壤肥、水条件对假臭草叶片的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积、气孔密度及气孔指数的影响显著。低光照及肥沃、湿润土壤生境与高光照及贫瘠、干旱土壤相比,假臭草的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积较大,气孔密度及气孔指数较小。【结论与意义】假臭草叶片气孔特征表现可塑性,说明其对异质环境具有一定的生态适应能力。     

Interpretation of an empirical model for stomatal conductance in terms of guard cell function

An empirical model for stomatal conductance (g), proposed by Leuning (1995, this issue) as a modification of Ball, Woodrow & Berry's (1987) model, is interpreted in terms of a simple, steady-state model of guard cell function. In this model, stomatal aperture is a function of the relative turgor between guard cells and epidermal cells. The correlation between g and leaf surface vapour pressure deficit in Leuning's model is interpreted in terms of stomatal sensing of the transpiration rate, via changes in the gradient of total water potential between guard cells and epidermal cells. The correlation between g, CO₂ assimilation rate and leaf surface CO₂ concentration in Leuning's model is interpreted as a relationship between the corresponding osmotic gradient, irradiance, temperature, intercellular CO₂ concentration and stomatal aperture itself. The explicit relationship between osmotic gradient and stomatal aperture (possibly describing the effect of changes in guard cell volume on the membrane permeability for ion transport) results in a decrease in the transpiration rate in sufficiently dry air. Possible extension of the guard cell model to include stomatal responses to soil water status is discussed.     

Modelling of stomatal density response to atmospheric CO2

Stomatal density tends to vary inversely with changes in atmospheric CO(2) concentration (C(a)). This phenomenon is of significance due to: (i) the current anthropogenic rise in C(a) and its impact on vegetation, and (ii) the potential applicability for reconstructing palaeoatmospheric C(a) by using fossil plant remains. It is generally assumed that the inverse change of stomatal density with C(a) represents an adaptation of epidermal gas conductance to varying C(a). Reconstruction of fossil C(a) by using stomatal density is usually based on empirical curves which are obtained by greenhouse experiments or the study of herbarium material. In this contribution, a model describing the stomatal density response to changes in C(a) is introduced. It is based on the diffusion of water vapour and CO(2), photosynthesis and an optimisation principle concerning gas exchange and water availability. The model considers both aspects of stomatal conductance: degree of stomatal aperture and stomatal density. It is shown that stomatal aperture and stomatal density response can be separated with stomatal aperture representing a short-term response and stomatal density a long-term response. The model also demonstrates how the stomatal density response to C(a) is modulated by environmental factors. This in turn implies that reliable reconstructions of ancient C(a) require additional information concerning temperature and humidity of the considered sites. Finally, a sensitivity analysis was carried out for the relationship between stomatal density and C(a) in order to identify critical parameters (= small parameter changes lead to significant changes of the results). Stomatal pore geometry (pore size and depth) represents a critical parameter. In palaeoclimatic studies, pore geometry should therefore also be considered.     

Optimal Control of Gas Exchange during Drought: Theoretical Analysis

An optimal strategy of stomatal control during a drought period,in plants adapted to a humid climate, is derived by maximizingthe photosynthetic production during the expected duration ofdrought. The expected duration of drought is calculated fromthe probability that rain occurs during a certain period, whichis assumed constant. The underlying plant model describes photosyntheticproduction and the consumption of water from the soil, witha given initial soil water content. Water is consumed throughtranspiration at a rate dependent on water vapour deficit, temperatureand stomatal conductance and carbon is assimilated at a ratedependent on light intensity and stomatal conductance. The optimizationproblem is solved with driving variables and the probabilityof rain corresponding to a Fenno-Scandian climate. The resultingoptimal stomatal control consists of two processes with differenttime constants: (1) daily variation depending on the drivingvariables, and (2) a declining trend as a function of the initialsoil water content and the probability of rain. The result allowsfor a physical interpretation of the so-called ‘cost ofwater’ used in similar optimization studies. An approximatemodel is derived from the optimal solution, such that the ‘costof water’ is a function of the soil water content. Photosynthesis; transpiration; stomatal conductance; soil water content; probability of rain; optimal control; drought; model     

Integration of hydraulic and chemical signalling in the control of stomatal conductance and water status of droughted plants

We describe here an integration of hydraulic and chemical signals which control stomatal conductance of plants in drying soil, and suggest that such a system is more likely than control based on chemical signals or water relations alone. The determination of xylem [ABA] and the stomatal response to xylem [ABA] are likely to involve the water flux through the plant. (1) If, as seems likely, the production of a chemical message depends on the root water status (Ψ_r), it will not depend solely on the soil water potential (Ψ_s) but also on the flux of water through the soil-plant-atmosphere continuum, to which are linked the difference between Ψ_r and Ψ_s. (2) The water flux will also dilute the concentration of the message in the xylem sap. (3) The stomatal sensitivity to the message is increased as leaf water potential falls. Stomatal conductance, which controls the water flux, therefore would be controlled by a water-flux-dependent message, with a water-flux-dependent sensitivity. In such a system, we have to consider a common regulation for stomatal conductance, leaf and root water potentials, water flux and concentration of ABA in the xylem. In order to test this possibility, we have combined equations which describe the generation and effects of chemical signals and classical equations of water flux. When the simulation was run for a variety of conditions, the solution suggested that such common regulation can operate. Simulations suggest that, as well as providing control of stomatal conductance, integration of chemical and hydraulic signalling may also provide a control of leaf water potential and of xylem [ABA], features which are apparent from our experimental data. We conclude that the root message would provide the plant with a means to sense the conditions of water extraction (soil water status and resisance to water flux) on a daily timescale, while the short-term plant response to this message would depend on the evaporative demand.     

Stomatal closure with soil water depletion not associated with changes in Bulk leaf water status

Summary It has previously been reported that canopy water loss by cowpea (Vigna unguiculata) decreases with small depletions in soil water. In these studies, under field conditions, it was demonstrated that with small changes in soil water status leaf conductance of cowpea decreases in a manner which is consistent with the sensitive regulation of canopy water loss.However, treatments which differed in leaf conductance, and presumably stomatal aperture, had similar leaf water potentials. It is hypothesized that the stomatal closure which results from soil water depletion is mediated by changes in root water status through effects on the flow of information from root to shoot. An efficient mechanism of this type could be partially responsible for the extreme drought avoidance exhibited by this plant.Dedicated to Dr. K. Springer     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.     

Comparing contributions of soil versus root colonization to variations in stomatal behavior and soil drying in mycorrhizal Sorghum bicolor and Cucurbita pepo

In prior studies we learned that colonization of soil can be as important as colonization of roots in determining mycorrhizal influence on the water relations of host plants. Here we use a path analysis modeling approach to test (a) whether quantity of hyphae in soil contributes to variations in stomatal behavior and soil drying, and (b) whether soil colonization or root colonization has a stronger influence on these stomatal and soil drying responses. Experiments were performed on Sorghum bicolor and Cucurbita pepo, with soils and roots colonized by a mixture of Glomus intraradices and Gigaspora margarita. Soil colonization generally made more significant contributions to stomatal conductance than did root colonization. Soil colonization did not make significant direct contributions to soil water potential measures (soil water potential at stomatal closure or soil drying rate), whereas root colonization did contribute a potentially important path to each. The findings further support a role for mycorrhization of the soil itself in contributing to the regulation of stomatal behavior of host plants.     

Evidence from Amazonian forests is consistent with isohydric control of leaf water potential

Climate modelling studies predict that the rain forests of the Eastern Amazon basin are likely to experience reductions in rainfall of up to 50% over the next 50-100 years. Efforts to predict the effects of changing climate, especially drought stress, on forest gas exchange are currently limited by uncertainty about the mechanism that controls stomatal closure in response to low soil moisture. At a through-fall exclusion experiment in Eastern Amazonia where water was experimentally excluded from the soil, we tested the hypothesis that plants are isohydric, that is, when water is scarce, the stomata act to prevent leaf water potential from dropping below a critical threshold level. We made diurnal measurements of leaf water potential (psi 1), stomatal conductance (g(s)), sap flow and stem water potential (psi stem) in the wet and dry seasons. We compared the data with the predictions of the soil-plant-atmosphere (SPA) model, which embeds the isohydric hypothesis within its stomatal conductance algorithm. The model inputs for meteorology, leaf area index (LAI), soil water potential and soil-to-leaf hydraulic resistance (R) were altered between seasons in accordance with measured values. No optimization parameters were used to adjust the model. This 'mechanistic' model of stomatal function was able to explain the individual tree-level seasonal changes in water relations (r2 = 0.85, 0.90 and 0.58 for psi 1, sap flow and g(s), respectively). The model indicated that the measured increase in R was the dominant cause of restricted water use during the dry season, resulting in a modelled restriction of sap flow four times greater than that caused by reduced soil water potential. Higher resistance during the dry season resulted from an increase in below-ground resistance (including root and soil-to-root resistance) to water flow.     

冬小麦近轴和远轴叶面气孔对土壤水分胁迫反应的敏感性

当根层土壤水分含量不足,作物体内出现水分胁迫时,小麦叶片两面气孔的反应有明显差异。远轴叶面气孔对水分胁迫的反应比近轴叶面气孔敏感。当出现水分胁迫时,远轴叶面气孔首先收缩,且收缩的程度比近轴叶面气孔大。远轴与近轴叶面气孔阻力的比值(r_(ab)/r_(ab))与根层平均土壤水势(Ψ_s)有关,当Ψ_s大于-50 kPa时,r_(ab)/r_(ad)基本稳定在1.5左右,而当Ψ_s小于-50 kPa时,r_(ab )/r_(ab)随Ψ_s降低而明显增加。     

The relations of stomatal closure and reopening to xylem ABA concentration and leaf water potential during soil drying and rewatering

Two tropical tree species, Acacia confusa and Leucaena leucocephala, were used to study the relationships among stomatal conductance, xylem ABA concentration and leaf water potential during a soil drying and rewatering cycle. Stomatal conductance of both A. confusa and L. leucocephala steadily decreased with the decreases in soil water content and pre-dawn leaf water potential. Upon rewatering, soil water content and pre-dawn leaf water potential rapidly returned to the control levels, whereas the reopening of stomata showed an obvious lag time. The length of this lag time was highly dependent not only upon the degree of water stress but also on plant species. The more severe the water stress, the longer the lag time. When A. confusa and L. leucocephala plants were exposed to the same degree of water stress (around –2.0 MPa in pre-dawn leaf water potential), the stomata of A. confusa reopened to the control level 6 days after rewatering. However, it took L. leucocephala about 14 days to reopen fully. A very similar response of leaf photosynthesis to soil water deficit was also observed for both species. Soil drying resulted in a significant increase in leaf and xylem ABA concentrations in both species. The more severe the water stress, the higher the leaf and xylem ABA concentrations. Both leaf ABA and xylem ABA returned to the control level following relief from water deficit and preceded the full recovery of stomata, suggesting that the lag phase of stomatal reopening was not controlled by leaf and/or xylem ABA. In contrast to drying the whole root system, drying half of the root system did not change the leaf water relations, but caused a significant increase in xylem ABA concentration, which could fully explain the decrease of stomatal conductance. After rewatering, the stomatal conductance of plants in which half of the roots were dried recovered more rapidly than those of whole-root dried plants, indicating that the leaf water deficit that occurred during the drying period was related to the post-stress stomatal inhibition. These results indicated that the decrease in stomatal conductance caused by water deficit was closely related to the increase in xylem ABA, but xylem ABA could not fully explain the reopening of stomata after relief of water stress, neither did the leaf ABA. Some unknown physiological and/or morphological processes in the guard cells may be related to the recovery process.     

Water deficits and hydraulic limits to leaf water supply

Many aspects of plant water use -- particularly in response to soil drought -- may have as their basis the alteration of hydraulic conductance from soil to canopy. The regulation of plant water potential (Psi) by stomatal control and leaf area adjustment may be necessary to maximize water uptake on the one hand, while avoiding loss of hydraulic contact with the soil water on the other. Modelling the changes in hydraulic conductance with pressure gradients in the continuum allows the prediction of water use as a function of soil environment and plant architectural and xylem traits. Large differences in water use between species can be attributed in part to differences in their 'hydraulic equipment' that is presumably optimized for drawing water from a particular temporal and spatial niche in the soil environment. A number of studies have identified hydraulic limits as the cause of partial or complete foliar dieback in response to drought. The interactions between root:shoot ratio, rooting depth, xylem properties, and soil properties in influencing the limits to canopy water supply can be used to predict which combinations should optimize water use in a given circumstance. The hydraulic approach can improve our understanding of the coupling of canopy processes to soil environment, and the adaptive significance of stomatal behaviour.     

The Simulation of Leaf Conductance Responses to Changes in Environmental Water Status

A simulation model of stomatal response to change of environmental water status was set up based on the works on the mechanism of stomatal movement. The variations of leaf conductance, water potential and turgot pressure in guard cells, subsidiary cells and the other cells or tissues in leaf with leaf-air vapour pressure difference and soil water potential have been calculated by our model. The calculated results fit very well with the data from experiments. The different patterns of leaf transpiration variation with the difference between leaf-air and vapour pressure can be explained quantitatively.     

冬小麦叶片气孔导度模型水分响应函数的参数化

植物气孔导度模型的水分响应函数用来模拟水分胁迫对气孔导度的影响过程, 是模拟缺水环境下植物与大气间水、碳交换过程的关键算法。水分响应函数包括空气湿度响应函数和土壤湿度(或植物水势)响应函数, 该研究基于田间实验观测, 分析了冬小麦(Triticum aestivum)叶片气孔导度对不同空气饱和差和不同土壤体积含水量或叶水势的响应规律。一个土壤水分梯度的田间处理在中国科学院禹城综合试验站实施, 不同水分胁迫下的冬小麦叶片气体交换过程和气孔导度以及其他的温湿度数据被观测, 同时观测了土壤含水量和叶水势。实验数据表明, 冬小麦叶片气孔导度对空气饱和差的响应呈现双曲线规律, 变化趋势显示大约1 kPa空气饱和差是一个有用的阈值, 在小于1 kPa时, 冬小麦气孔导度对空气饱和差变化反应敏感, 而大于1 kPa后则反应缓慢; 分析土壤体积含水量与中午叶片气孔导度的关系发现, 中午叶片气孔导度随土壤含水量增加大致呈现线性增加趋势, 但在平均土壤体积含水量大于大约25%以后, 气孔导度不再明显增加, 而是维持在较高导度值上下波动; 冬小麦中午叶片水势与相应的气孔导度之间, 随着叶水势的增加, 气孔导度呈现增加趋势。根据冬小麦气孔导度对空气湿度、土壤湿度和叶水势的响应规律, 研究分别采用双曲线和幂指数形式拟合了水汽响应函数, 用三段线性方程拟合了土壤湿度响应函数和植物水势响应函数, 得到的参数可以为模型模拟冬小麦的各类水、热、碳交换过程采用。     

Stomatal behaviour and leaf water potential of chilled and water-stressed Solanum melongena, as influenced by growth history

Abstract Leaf diffusion resistance and leaf water potential of intact Solanum melongena plants were measured during a period of chilling at 6 °C. Two pretreatments, consisting of a period of water stress or a foliar spraying of abscisic acid (ABA), were imposed upon the plants prior to chilling. The control plants did not receive a pretreatment. In addition to intact plant studies, stomatal responses to water loss and exogenous abscisic acid were investigated using excised leaves, and the influence of the pretreatment observed. Chilled, control plants wilted slowly and maintained open stomata despite a decline in leaf water potential to –2.2 MPa after 2 d of chilling. In contrast plants that had been water stressed or had been sprayed with abscisic acid, prior to chilling, did not wilt and maintained a higher leaf water potential and a greater leaf diffusion resistance. In plants that had not received a pretreatment, abscisic acid caused stomatal closure at 35 °C, but at 6°C it did not influence stomatal aperture. The two pretreatments greatly increased stomatal sensitivity to both exogenous ABA and water stress, at both temperatures. Stomatal response to water loss from excised leaves was greatly reduced at 6°C. These results are discussed in relation to low temperature effects on stomata and the influence of preconditioning upon plant water relations.     

玉米叶片气孔及花环和维管束结构对水分胁迫的响应

采用盆栽种植,以玉米品种郑单958为试验材料,设置对照（CK）、轻度（LS）、中度（MS）和重度（SS）水分胁迫 (土壤含水量分别为田间持水量的75%～85%、65%～75%、55%～65%、45%～55%)4个水分梯度,从气孔开度的调控、花环结构的变化、叶片维管束水分运输等方面研究了玉米对土壤水分胁迫的应激反应.结果表明：随着水分胁迫程度的不断加剧,气孔保卫和副卫细胞中过氧化氢(H₂O₂)的积累量逐渐增多,应用荧光染色定位也发现H₂O₂荧光强度逐渐增强,而气孔开度和气孔导度均逐渐减小.同时,花环的正常结构被破坏,花环细胞排列凌乱且体积逐渐变小,维管束鞘细胞变得不规则;大维管束断面面积、木质部面积以及韧皮部细胞数均减少,总的叶片和上、下表皮的厚度逐渐变薄.此外,花环细胞和维管束鞘细胞中叶绿体数目减少,且在中度胁迫下花环细胞中叶绿体的分布发生了变化,由紧贴细胞质膜内侧环靠细胞壁分布向偏细胞中心扩散.发现玉米气孔关闭可能是由保卫细胞和副卫细胞中的H₂O₂共同调节,副卫细胞中的H₂O₂对保卫细胞主导的气孔关闭具有协同作用.总之,在水分胁迫下,玉米通过改变叶片花环结构和厚度、叶绿体的分布,减小木质部和韧皮部面积等降低叶片表面水势,促进气孔关闭,减少体内水分散失,以减轻干旱胁迫对其伤害.     

Use of the root contact concept,an empirical leaf conductance model and pressure-volume curves in simulating crop water relations

A simulation model “DanStress” was developed for studying the integrated effects of soil, crop and climatic conditions on water relations and water use of field grown cereal crops. The root zone was separated into 0.1 m deep layers of topsoil and subsoil. For each layer the water potential at the root surface was calculated by a single root model, and the uptake of water across the root was calculated by a root contact model. Crop transpiration was calculated by Monteith's combination equation for vapour flow. Crop conductance to water vapour transfer for use in Monteith's combination equation was scaled up from an empirical stomatal conductance model used on sunlit and shaded crop surfaces of different crop layers. In the model, transpirational water loss originates from root water uptake and changes in crop water storage. Crop water capacitance, used for describing the water storage, was derived from the slope of pressure-volume (PV) curves of the leaves. PV curves were also used for deriving crop water potential, osmotic potential, and turgor pressure. The model could simulate detailed diurnal soil-crop water relations during a 23-day-drying cycle with time steps of one hour. During the grain filling period in spring barley (Hordeum distichum L.), grown in a sandy soil in the field, measured and predicted values of leaf water and osmotic potential, RWC, and leaf stomatal conductance were compared. Good agreement was obtained between measured and predicted values at different soil water deficits and climatic conditions. In the field, measured and predicted volumetric soil water contents (θ) of topsoil and subsoil layers were also compared during a drying cycle. Predicted and measured θ-values as a function of soil water deficits were similar suggesting that the root contact model approach was valid. From the investigation we concluded: (I) a model, which takes the degree of contact between root surface and soil water into account, can be used in sandy soil for calculation of root water uptake, so that the root conductance during soil water depletion only varies by the degree of contact; (II) crop conductance, used for calculation of crop transpiration, can be scaled up from an empirical single leaf stomatal conductance model controlled by the level of leaf water potential and micrometeorological conditions; (III) PV curves are usable for describing crop water status including crop water storage.     

Stomatal control of transpiration in the canopy of a clonal Eucalyptus grandis plantation

 Predawn leaf water potential, stomatal conductance and microclimatic variables were measured on 13 sampling days from November 1995 through August 1996 to determine how environmental and physiological factors affect water use at the canopy scale in a plantation of mature clonal Eucalyptus grandis Hill ex-Maiden hybrids in the State of Espirito Santo, Brazil. The simple ”big leaf” Penman-Monteith model was used to estimate canopy transpiration. During the study period the predawn leaf water potential varied from –0.4 to –1.3 MPa, with the minimum values observed in the winter months (June and August 1996), while the average estimated values for canopy conductance and canopy transpiration fell from 17.3 to 5.8 mm s^–1 and from 0.54 to 0.18 mm h^–1, respectively. On the basis of all measurements, the average value of the decoupling coefficient was 0.25. During continuous soil water shortage a proportional reduction was observed in predawn leaf water potential and in daily maximum values of stomatal conductance, canopy transpiration and decoupling coefficient. The results showed that water vapour exchange in this canopy is strongly dominated by the regional vapour pressure deficit and that canopy transpiration is controlled mainly by stomatal conductance. On a seasonal basis, stomatal conductance and canopy transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture and rainfall. Good results were obtained with a multiplicative empirical model that uses values of photosynthetically active radiation, vapour pressure deficit and predawn leaf water potential to estimate stomatal conductance. Received: 10 June 1998 / Accepted: 20 July 1998