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.     

马占相思林冠层气孔导度对环境驱动因子的响应

利用Granier热消散探针在2003年10月测定了广东鹤山丘陵地马占相思林14株样树的树干液流,同时监测林冠上方的光合有效辐射、空气湿度和气温,结合树木的形态和林分的结构特征,计算马占相思的整树蒸腾(E)、林分总蒸腾(E_t)以及冠层平均气孔导度(g_c),分析树形特征与整树水分利用的关系、冠层气孔导度对光合有效辐射(PAR)和空气水汽压亏缺(D)的响应.结果表明,整树蒸腾与胸径(P＜0.0001)、边材面积(P＜0.0001)和冠幅(P=0.0007)以自然对数的形式、与树高(P=0.014)以幂函数的形式呈现显著正相关.冠层气孔导度最大值(g_cmax)随D的上升呈对数函数下降(P＜0.0001),对光合有效辐射的响应则呈双曲线函数增加(P＜0.0001).液流测定系统能提供连续和准确的整树和林分蒸腾速率值,经严格数学推导公式计算,最终可求出冠层气孔导度,是研究森林水分利用与环境因子相互关系的有效方法.     

A model for simulating transpiration of Eucalyptus salmonophloia trees

Understanding the water relations of Eucalyptus trees plays an important role in finding solutions to dryland salinity in southern Australia. A model for studying structure–function relationships in isolated tree crowns (radiation absorption, transpiration and photosynthesis, RATP) was parameterized to permit the seasonal transpiration course of a Eucalyptus salmonophloia tree to be quantified. Model responses to different parameterizations were tested in a sensitivity analysis. Predictive quality was mostly affected by the accuracy of information about leaf area density and stomatal responses to air vapor pressure deficit, and to a lesser extend by foliage dispersion. Assuming simple, non‐synergistic influences of changes in photosynthetic active radiation and air vapor pressure deficit on stomatal transpiration control, the model was able to simulate the daily water uptake of E. salmonophloia trees with reasonable predictive quality during an entire season. In order to more precisely simulate short‐term (i.e. diurnal) water use dynamics, the model must be extended to account for hydraulic and chemical controls of stomatal regulation of crown energy balance.     

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     

不同水分条件下盆栽苹果树蒸腾速率动态模拟

土壤水分是制约作物产量和品质的主要环境因素之一,估算不同水分条件下的蒸腾速率（r3对于作物的优质高产和节水灌溉等具有十分重要的意义。通过构建蒸腾．气孔．光合耦合模型可模拟出不同水分条件下苹果树的蒸腾动态,模型参数根据逐步干旱条件下盆栽‘富士’苹果树试验获取。结果表明,Tr主要由饱合水气压差和气孔导度（G。）驱动,同时气象因子和土壤水势对其有强烈的交互作用影响。Tr随土壤水势的下降而减小,当土壤水势低于-0．4MPa时减小幅度更加显著。晴天时,G。在一天中呈双峰曲线,而一呈单峰曲线,最大值出现在13：00左右,约为3．6mmol·m-Ls。根据该模型可计算出不同水分条件下1株盆栽苹果树（总叶片积为O．26m-2）全天的蒸腾总量,供水充足时为652．1g,严重干旱时（土壤水势为-1．5MPa）为85．4g。实测值和模拟值的比较表明,该耦合模型能够模拟出不同土壤水分条件下盆栽苹果树的蒸腾动态以及土壤的含水量。     

Transpiration from a mature Eucalyptus globulus plantation in Portugal during a spring-summer period of progressively higher water deficit

The rates of transpiration from a mature Eucalyptus globulus Labill. stand in Portugal were evaluated during a drying period of the spring-summer 1994. Transpiration was measured by the Granier sap flow method and estimated by the Penman-Monteith model. During the experimental period daily transpiration varied between 3.64 and 0.50 mm day⁻¹. For high-transpiration days, a good agreement was observed between Penman-Monteith estimates and sap flow measurements, both on a daily and on an hourly basis. However, for low-transpiration days, the Penman-Monteith model overestimated transpiration in comparison with the sap flow method. The diurnal variation of sap flow was then smoother and lagged behind the estimates of the Penman-Monteith model. E. globulus showed an efficient control of transpiration losses during dry periods through a progressive stomatal closure. As soil moisture deficit increased, the daily maximum stomatal conductance decreased from 0.46 to 0.14 cm s⁻¹. The results also show that, on a seasonal basis, stomatal conductance and daily transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture content. Received: 26 January 1996 / Accepted: 20 October 1996     

Differences in transpiration of Norway spruce drought stressed trees and trees well supplied with water

The paper focuses on the evaluation of transpiration as a physiological process, which is very sensitive to drought stress. Reactions of 25-year-old Norway spruce (Picea abies (L.) Karst.) trees to drought were examined during 2009 summer. Sap flow rate (SF), meteorological and soil characteristics were measured continually. Vapour pressure deficit of the air (VPD) and cumulative transpiration deficit (KTD) was calculated. During the second half of the vegetation period, the decrease in soil water content was observed and irrigation was applied to a group of spruce trees, while the second group was treated under natural soil drought. On the days, when the differences in transpiration between irrigated (IR) and non-irrigated (NIR) trees were significant (21 days), transpiration of NIR trees was only 23% of the transpiration of IR trees. We found significant differences in transpiration when the soil water content (SWC) of NIR variant at a depth of 5–15 cm ranged from 10.4 to 13.7%. Under both regimes of water availability, daily transpiration significantly responded to atmospheric conditions. However, the influence of all assessed meteorological parameters on SF of NIR trees was significantly lower than on IR tree. The dependency of transpiration on evaporative demands of atmosphere decreased with the decreasing soil moisture. Cumulative transpiration deficit of the stand during the entire evaluated period was 50.9 mm. The difference between the transpiration of the mean NIR tree and of the mean IR tree was 278.8 L over the assessed period of 47 days (5.9 L per day). The transpiration of NIR trees was 40.3% from the transpiration of IR trees during this period.     

Transpiration and visual appearance of warm season turfgrasses during soil drying

Warm-season turfgrasses may be subjected to increasing drought as future urban irrigation regulations become more restrictive. Species differences in water use and transpiration response to drying soil may be exploited in the future to increase survival and maintain green color under drying soil conditions. This study was undertaken to provide background documentation on the sensitivity to soil–water deficit of three warm-season grasses: ‘Argentine’ bahiagrass (Paspalum notatum); ‘Floratam’ St. Augustinegrass (Stenotaphrum secundatum), and ‘Empire’ zoysiagrass (Zoysia japonica). Each of these turfgrasses demonstrated a two-phased linear transpiration response to gradually drying soil as expressed by a normalized ratio between the transpiration rates of drought stressed to well-watered plants (NTR). In this study, well-watered bahiagrass used 30% more water on a daily basis than did well-watered St. Augustinegrass or zoysiagrass. However, under drought, the three grass species transpired the same amount of water during the soil drying period up until NTR to 0.1. Since bahiagrass reached an NTR of 0.1 at 10.3 days versus 12.7 and 13.0 days for St. Augustinegrass and zoysiagrass, respectively, bahiagrass demonstrated a more rapid water loss rate during the drying period. The fraction of transpirable soil water (FTSW) remaining in the soil at the breakpoints for bahiagrass, St. Augustinegrass and zoysiagrass were 0.13, 0.16, and 0.19, respectively, in 2010, but were 0.18, 0.30, and 0.22, respectively, under slightly warmer conditions in 2011. The consistently low FTSW breakpoint for bahiagrass means that compared to the other species, bahiagrass continues to use water at a high rate late into the soil drying cycle before conserving soil water by decreasing stomatal conductance. That is, bahiagrass is likely to be subjected to greater soil–water deficits in lengthy droughts and needs mechanisms to better survive these droughts. The differences in breakpoints by year may be due to a combination of soil factors and temperature differences in the greenhouse.     

Irrigation effects on daily and seasonal variations of trunk sap flow and leaf water relations in olive trees

Irrigation effects on whole-plant sap flow and leaf-level water relations were characterised throughout a growing season in an experimental olive (Olea europaea L.) orchard. Atmospheric evaporative demand and soil moisture conditions for irrigated and non-irrigated olive trees were also monitored. Whole-plant water use in field-grown irrigated and rain fed olive trees was determined using a xylem sap flow method (compensation heat-pulse velocity). Foliage gas exchange and water potentials were determined throughout the experimental period. Physiological parameters responded diurnally and seasonally to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the Penman–Monteith equation in the field. Summer drought caused decreasing leaf gas exchange and water potentials, and a progressive increase in hydraulic conductance (stronger in non-irrigated than irrigated trees), probably attributable to modifications in hydraulic properties at the soil-root interface. Negligible hysteresis, attributable to low plant capacitance, was observed in the relationship between leaf water potential and sap flow. A proportional decrease in maximum daily leaf conductance with increasing vapour pressure deficit was observed, while mean daytime canopy stomatal conductance decreased with the season. As a result, plant water use was limited and excessive drought stress prevented. Non-irrigated olive trees recovered after the summer drought, showing a physiological behaviour similar to that of irrigated trees. In addition to physiological and environmental factors, there are endogenous keys (chemical signals) influencing leaf level parameters. Olive trees are confirmed to be economical and sparing users of soil water, with an efficient xylem sap transport, maintenance of significant gas exchange and transpiration, even during drought stress.     

Effects of soil water deficit and nitrogen nutrition on water relations and photosynthesis of pot-grown Coffea canephora Pierre

Coffea canephora plants (clone INCAPER-99) were submitted to low N (LN) or high N (HN) applications and two watering regimes (daily irrigation and irrigation every 5 days for a month). Although water potential was not altered significantly by N, HN plants showed higher relative water content than did LN plants under water deficit. Only HN plants exhibited some ability for osmotic adjustment. Plants from both N treatments increased their cell wall rigidity under drought, with a more pronounced augmentation in HN plants. In well-watered plants, carbon assimilation rate increased with increasing N while stomatal conductance did not respond to N supply. Under drought conditions, carbon assimilation decreased by 68-80% compared to well-watered plants, whereas stomatal conductance and transpiration rate declined by 35% irrespective of the N applications. Stable carbon isotope analysis, combined with leaf gas exchange measurements, indicated that regardless of the watering treatments, N increased the long-term water use efficiency through changes in carbon assimilation with little or no effect on stomatal behaviour.     

The photosynthetic response of tobacco plants overexpressing ice plant aquaporin McMIPB to a soil water deficit and high vapor pressure deficit

We investigated the photosynthetic capacity and plant growth of tobacco plants overexpressing ice plant (Mesembryanthemum crystallinum L.) aquaporin McMIPB under (1) a well-watered growth condition, (2) a well-watered and temporal higher vapor pressure deficit (VPD) condition, and (3) a soil water deficit growth condition to investigate the effect of McMIPB on photosynthetic responses under moderate soil and atmospheric humidity and water deficit conditions. Transgenic plants showed a significantly higher photosynthesis rate (by 48 %), higher mesophyll conductance (by 52 %), and enhanced growth under the well-watered growth condition than those of control plants. Decreases in the photosynthesis rate and stomatal conductance from ambient to higher VPD were slightly higher in transgenic plants than those in control plants. When plants were grown under the soil water deficit condition, decreases in the photosynthesis rate and stomatal conductance were less significant in transgenic plants than those in control plants. McMIPB is likely to work as a CO₂ transporter, as well as control the regulation of stomata to water deficits.     

基于树干液流测定值进行尺度扩展的马占相思林段蒸腾和冠层气孔导度

应用Granier热消散探针,长期监测华南丘陵地马占相思(Acacia mangium)林14棵样树的树干液流(Sap flow),由此计算整树和林段的蒸腾速率,结合同步记录的环境因子,求算冠层平均气孔导度(G_c)。Granier探针的灵敏度较高,能精确测定即使是微弱的液流活动。观测结果显示,树木个体之间的液流密度(J_s)和整树蒸腾(E_t)受树形特征影响较大。马占相思林径级大的树木个体数较少,但占据林段边材总面积和林段蒸腾的比例较大。J_s和E_t的日变化主要受光合有效辐射(Q_o)和空气水蒸气压亏缺(D)的控制,土壤含水量(θ)对较大胸径树木E_t的影响大于胸径较小的树木,个体之间J_s和E_t的差异随θ的下降而缩小。一年中,林段蒸腾(E)在光照和水热条件较好的7月最高,9~12月,由于土壤水分供应的减少致使E值降低,E对D的敏感性下降。G_c与主要环境因子的关系与E相似,如果θ长期偏低,G_c会明显下降,是造成E降低的主要原因。成熟马占相思林在光照充足、水热条件较好的情况下的蒸腾活动旺盛,但对土壤水分胁迫的忍受力较低。     

Effects of Di-n-Butyl Phthalate on Mycorrhizal and Non-Mycorrhizal Cowpea Plants

This study examined the effect of mycorrhizal colonization with Glomus intraradices on physiological parameters and foliar nutrient concentrations in Olea europaea L. subsp. sylvestris and Rhamnus lycioides L. seedlings subjected to well-watered or drought-stressed conditions. Under drought stress, mycorrhizal O. europaea seedlings showed significantly higher photosynthetic and transpiration rates, stomatal conductance and foliar P concentration than its similarly-sized non-mycorrhizal counterpart. The intrinsic water use efficiency (photosynthetic rate to stomatal conductance ratio) was not change in O. europaea and decreased in R. lycioides seedlings due to mycorrhizal colonization under both well-watered and drought-stress conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

空气湿度与土壤水分胁迫对紫花苜蓿叶表皮蜡质特性的影响

选用2个抗旱性不同的紫花苜蓿品种,敖汉(强抗旱)和三得利(弱抗旱),设置空气湿度(45%-55%和75%-85%)和土壤水分胁迫(75%和35%田间持水量)处理,分析紫花苜蓿叶表皮蜡质含量、组分及晶体结构、气体交换参数、水势及脯氨酸含量的变化规律。结果表明,单独土壤水分胁迫时,紫花苜蓿叶表皮蜡质晶体结构及蜡质总量无显著变化;敖汉蜡质组分中烷类、酯类含量增加,醇类含量下降;三得利醇类含量下降,烷类、酯类含量变化不显著。低空气湿度胁迫时,两品种蜡质总量无显著变化,烷类和酯类含量显著增加,醇类含量显著下降,叶表皮片状蜡质晶体结构熔融呈弥漫性,扩大了对叶表面积的覆盖,其蒸腾速率显著低于正常湿度。复合胁迫处理时,叶表皮片状蜡质晶体结构继续呈弥漫性,烷类、酯类、未知蜡质组分含量均高于单独胁迫处理,醇类含量最低,而蜡质总量除三得利显著高于对照外,其余均无显著差异。紫花苜蓿叶表皮蜡质各组分含量(除醇类)及蜡质总量与光合速率呈显著负相关,与蒸腾速率无显著相关关系。蜡质总量与叶水势呈显著正相关。总体上,敖汉蜡质总量显著高于三得利,蜡质组分中烷类物质的增加有助于提高植株的抗旱性。在复合胁迫下,强抗旱品种主要通过气孔因素控制水分散失,而弱抗旱品种通过气孔和非气孔因素共同控制植物水分散失。     

Effects of drought-rewatering-drought on photosynthesis and growth of maize

Aims Our objective was to investigate the responses of maize photosynthesis and growth to repeated drought.Methods Maize seedlings were exposed to different soil water deficit for three weeks, then rewatering for one week, and again to different water deficit for three weeks, to examine the effects of repeated drought on photosynthesis and growth.Important findings After the first water deficit treatments, under severe drought, plant height, total leaf area of individual plant, shoot and root biomass declined significantly, also transpiration rate (T_r), stomatal conductance (G_s), intercellular CO₂ concentration (C_i), net photosynthetic rate (P_n), maximum net photosynthetic rate (A_max), but light compensation point and dark respiration rate increased significantly. Under medium drought, plant height, leaf area, and shoot biomass decreased significantly, but root biomass did not vary, hence, the ratio of roots to shoots (R/S) increased. Moreover, plants did not show significant differences in photosynthetic parameters. After rewatering, photosynthesis and growth rate of plants previously exposed to water deficit could recover to the levels of well-watered plants, but plant height and leaf area did not recover to the levels of the control. When maize were subjected to recurrent drought, plants pre-exposed to medium drought showed no significant difference in plant height, biomass, and photosynthetic parameters, but a significant decrease in leaf area, compared to plants only exposed to second medium drought. Plants pre-exposed to severe drought had significantly higher T_r, G_s, C_i, P_n, A_max, and, apparent quantum yield but significantly lower plant height, leaf area, and biomass than plants without previous exposure. These results indicated that the first severe drought significantly reduced photosynthetic capacity and maize growth, rewatering could recover photosynthesis and growth rate to the levels of well-watered plants, but could not eliminate the adverse influence of the first drought on growth. The first medium drought could stimulate the growth of maize root system and significantly increased R/S, which can enhance maize drought resistance to subsequent repeated drought, and maintain the total biomass in the control level; the first severe drought could enhance maize drought resistance to subsequent repeated drought in the aspect of photosynthesis, but could not compensate for the adverse effect of early drought on plant growth. Hence, in practice, drought hardening should be limited in the level of medium drought, and avoiding severe drought.     

干旱-复水-再干旱处理对玉米光合能力和生长的影响

为了探求玉米(Zea mays)光合作用和生长对重复干旱的响应机制, 采用盆栽试验, 分别测定了不同程度土壤干旱处理3周时、随后复水1周时以及再次不同程度干旱处理3周时玉米幼苗光合参数和生长的变化。第一次土壤干旱处理后, 重度干旱处理显著降低玉米株高、单株总叶面积、地上部分及根系生物量以及叶片的蒸腾速率(T_r)、气孔导度(G_s)、胞间CO₂浓度(C_i)、净光合速率(P_n)和最大净光合速率(A_max), 但显著提高光补偿点和暗呼吸速率; 中度干旱处理同样显著降低玉米株高、叶面积和地上部分生物量, 但对根系生物量无影响, 因而根冠比增大, 对上述光合参数的负效应也不具有显著性。复水可使前期经受中度和重度干旱处理的玉米植株的光合能力和生长速率恢复到正常水分条件下生长的植株的水平, 但株高和叶面积没有恢复到对照水平。当玉米再次经受水分亏缺处理时, 与只遭受第二次中度或重度干旱处理的植株相比, 经历过前期中度干旱处理的植株的株高、生物量和光合参数没有显著变化, 但叶面积显著下降; 经历过前期重度干旱处理植株的T_r、G_s、C_i、P_n、A_max和表观量子效率显著升高, 而株高、叶面积和生物量显著降低。综上所述, 第一次重度干旱处理显著降低玉米叶片的光合能力和生长, 复水可使光合能力和生长速率恢复到正常水分条件下生长植株的水平, 但不能消除前期干旱对生长产生的不利影响。前期中度干旱可以刺激玉米根系的生长和显著提高根冠比, 有利于提高对二次干旱的抵抗能力, 并使总的生物量保持在对照水平, 而前期重度干旱处理虽然在光合作用上能提高植株对二次干旱的抵御能力, 但不能弥补前期干旱处理对生长的不利影响。因此, 在生产实践中, 如果进行抗旱锻炼, 应限制在中度干旱水平, 避免重度干旱。     

Long-term transpiration change with rainfall decline in a Mediterranean Quercus ilex forest

In the Mediterranean basin, precipitation is expected to decline as a consequence of climate change. The response of a Quercus ilex forest in southern France to such a decline in water availability was studied using a 4-year throughfall exclusion experiment. Seasonal courses of sap flow and leaf water potential were obtained from 2004 to 2007 and used to characterize tree water relations in a control and a dry treatment. The experiment reduced the average precipitation input to the soil by 29%, and resulted in a 23% reduction in annual transpiration. Soil water potential was significantly lower in the dry treatment only during summer drought, but transpiration was reduced all year round even during well-watered periods. Despite a tight stomatal control over transpiration, whole-tree hydraulic conductance was found to be lower in the trees growing in the driest conditions. This reduction in water transport capacity was observed jointly with a reduction in leaf transpiring area. Canopy leaf area decreased by 18% in the dry treatment as a consequence of the throughfall exclusion, which was found to validate the ecohydrological equilibrium theory.     

Influence of soil,plant and meteorological factors on water relations and yield inHevea brasiliensis

Influence of factors governing the soil-plantatmosphere system on components of water relations and yield was studied in two clones of rubber tree,Hevea brasiliensis, viz. RRII 105 and RRII 118. Clonal variations were evident in yield and yield components and associated physiological parameters in response to soil moisture status and meteorological factors. Observations made during different seasons indicatedvariations in yield are attributed to differences in plugging index and initial flow rates, to the major yield components and also variations in components of water relations as influenced by meteorological factors. Among the two clones, RRII 105 was found to be fairly drought tolerant compared to RRII 118. RRII 105 was found to respond well to dry weather through higher stomatal resistances, higher leaf water potentials, lowered transpirational water loss and lower relative transpiration ratios, while RRII 118 was susceptible to stress situations.     

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