Effects of Atmospheric Saturation Deficit on the Stomatal Conductance of Pearl Millet (Pennisetum typhoides S. and H.) and Groundnut (Arachis hypogaea L.)

Experiments were carried out to investigate the influence ofatmospheric saturation deficit on the stomatal conductancesof millet and groundnut plants grown in undisturbed soil incontrolled environment glasshouses. Environmental conditionsduring growth were maintained close to those experienced inthe semi-arid tropics. The results demonstrated that the stomatal conductances of well-wateredplants of both species were affected strongly by changes insaturation deficit. The response was stronger at higher irradianceswhen variations in saturation deficit between 1.5 and 3.0 kPacaused 3–4-fold changes in leaf conductance. However,the stomatal response was greatly reduced or absent in unirrigatedplants in which stomatal conductances were reduced. Reduction of the transpiring leaf area by covering some of theleaves increased the leaf conductances of the remaining leavesand partially restored the stomatal response to saturation deficitin unirrigated plants. Leaf conductance was sensitive to thetranspiring area per plant and declined as the transpiring areaincreased. However, the reduction in mean leaf conductance wasinsufficient to prevent an increase in canopy conductance owingto the increased transpiring area: The results are compared to earlier field data for millet, andthe possible origin of the stomatal response is discussed.     

植物气孔扩散传导率的研究

本文对生长在可控环境温室中的花生气孔扩散传导率进行了实验研究,揭示了单个植株之间、上下表面之间、叶片不同部位以及冠层垂直方向上气孔扩散传导率的变异性。同时以气孔扩散传导率与环境条件的测定值为基础,对传导率对环境因子的反应进行了分析,植株顶部叶片气孔扩散传导率与太阳总辐射和空气饱和差有关系;冠层传导率与冠层截留辐射和空气饱和差有相关关系。     

Stomatal control of transpiration from a developing sugarcane canopy

Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.     

Combining thermal and visible imagery for estimating canopy temperature and identifying plant stress

Thermal imaging is a potential tool for estimating plant temperature, which can be used as an indicator of stomatal closure and water deficit stress. In this study, a new method for processing and analysing thermal images was developed. By using remote sensing software, the information from thermal and visible images was combined, the images were classified to identify leaf area and sunlit and shaded parts of the canopy, and the temperature statistics for specific canopy components were calculated. The method was applied to data from a greenhouse water-stress experiment of Vicia faba L. and to field data for Vitis vinifera L. Vaseline-covered and water-sprayed plants were used as dry and wet references, respectively, and two thermal indices, based on temperature differences between the canopy and reference surfaces, were calculated for single Vicia faba plants. The thermal indices were compared with measured stomatal conductance. The temperature distributions of sunlit and shaded leaf area of Vitis vinifera canopies from natural rainfall and irrigation treatments were compared. The present method provides two major improvements compared with earlier methods for calculating thermal indices. First, it allows more accurate estimation of the indices, which are consequently more closely related to stomatal conductance. Second, it gives more accurate estimates of the temperature distribution of the shaded and sunlit parts of canopy, and, unlike the earlier methods, makes it possible to quantify the relationship between temperature variation and stomatal conductance.     

Stomatal responses of pearl millet (Pennisetum americanum [L.] Leeke) to leaf water status and environmental factors in the field

Abstract. Factors affecting stomatal conductance (g₁) of pearl millet ( Pennisetum americanum [L.] Leeke), cultivar BJ 104, were examined in the field in India during the dry season.
Diurnal changes in g₁ were evaluated for upper expanded leaves at flowering on two occasions using plants subjected to varying degrees of water stress. Except for the most severely stressed treatment, diurnal changes in g₁ closely matched changes in irradiance ( I ), the promotive effect of which largely overcame opposing influences on g₁ of increasing atmospheric vapour pressure deficit, and decreasing leaf water and turgor potentials (Ψ, Ψ_p).
Two main effects of water stress on g₁ were evident: (i) a decrease in the amplitude of the mid-day peak in g₁, and (ii) a decrease in the time over which high g₁ was maintained, resulting in early (mid-day) closure and hysteresis in the relationship between g₁ and I .
Leaf conductance was greatest for upper leaves and decreased down the canopy. At equivalent depths in the canopy g₁ was higher in flowering than in photoperiodically-retarded plants of the same age. The magnitude of water stress-induced stomatal closure increased down the plant, and was more marked in retarded than in flowering plants.
Within individual stress treatments Ψ of upper leaves decreased linearly as transpiration flux increased. It is concluded that stomatal behaviour of upper leaves of pearl millet at flowering largely operates to maximize assimilation rather than to minimize water loss.     

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     

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。     

Stomatal responses to increased CO2: implications from the plant to the global scale

Increased atmospheric CO₂ often but not always leads to large decreases in leaf conductance. Decreased leaf conductance has important implications for a number of components of CO₂ responses, from the plant to the global scale. All of the factors that are sensitive to a change in soil moisture, either amount or timing, may be affected by increased CO₂. The list of potentially sensitive processes includes soil evaporation, run-off, decomposition, and physiological adjustments of plants, as well as factors such as canopy development and the composition of the plant and microbial communities. Experimental evidence concerning ecosystem-scale consequences of the effects of CO₂ on water use is only beginning to accumulate, but the initial indication is that, in water-limited areas, the effects of CO₂-induced changes in leaf conductance are comparable in importance to those of CO,₂-induced changes in photosynthesis. Above the leaf scale, a number of processes interact to modulate the response of canopy or regional evapotran-spiration to increased CO₂. While some components of these processes tend to amplify the sensitivity of evapo-transpiration to altered leaf conductance, the most likely overall pattern is one in which the responses of canopy and regional evapotranspiration are substantially smaller than the responses of canopy conductance. The effects of increased CO₂ on canopy evapotranspiration are likely to be smallest in aerodynamically smooth canopies with high leaf conductances. Under these circumstances, which are largely restricted to agriculture, decreases in evapotranspiration may be only one-fourth as large as decreases in canopy conductance. Decreased canopy conductances over large regions may lead to altered climate, including increased temperature and decreased precipitation. The simulation experiments to date predict small effects globally, but these could be important regionally, especially in combination with radiative (greenhouse) effects of increased CO₂.     

Applying hydraulic lift in an agroecosystem: forage plants with shoots removed supply water to neighboring vegetable crops

When a plant encounters spatially heterogeneous soil moisture within its root system, usually drier surface and moister subsurface soils, water can move between these layers through the root system, a plant process known as hydraulic lift or redistribution. The water thus transferred is available not only for the plant itself but also for its neighbors. We examined application of this process as a possible biological irrigation tool. As ??donors??, we used perennial forage plants with their shoots removed to minimize the effect of light-interception by them on the ??receiver?? plants growing alongside them. In a horizontally split-root experiment, where an upper container was filled with sand and a lower one with water, superior donor species could maintain the upper sand in a fully hydrated condition for several weeks, increasing stomatal conductance in the receivers. The effects were also confirmed in a water-limited agricultural field, as significant differences were found in canopy temperature and yield in neighboring crop plants in the presence or absence of donor root systems. These results suggest that deep-rooting associate plants with their shoots removed function as an irrigation tool and improve crop production in water-scarce environments.     

Modelling the surface conductance of a broad-leaf canopy: effects of partial decoupling from the atmosphere

The transpiration of a mature beech ( Fagus sylvatica L.) forest was measured over a whole season by the heat pulse velocity technique and the results analysed in terms of a new analytical canopy conductance model, which takes into account the effects of partial decoupling from the atmosphere on the local humidity environment experienced by the canopy. Stand daily transpiration ranged from 0·62 to 2·97 mm d^–1, with a seasonal mean value of 1·97 mm d^–1. Maximum canopy conductance was 18·5 mm s^–1, with a mean estimated value of 5·0 mm s^–1; computed values were little affected by the assumption of neutral atmospheric conditions. The decoupling coefficient Ω varied greatly on a daily and seasonal basis, with an average value of 0·28. As a result of partial decoupling, the estimated vapour pressure deficit at the notional canopy surface exceeded the values measured above the canopy by 380 Pa on average. When correctly expressed in terms of humidity at the canopy surface, the model explained 80% of the variance in half-hourly transpiration measurements. Upon cross-validation it still explained 72% of the variance, as compared to only 40% when correction for partial decoupling was not introduced. A baseline canopy conductance of 0·7 mm s^–1, not modulated by the environment, was estimated. The implications of the model are discussed for the representation of canopy conductance and transpiration of broad-leaf forests.     

Evaluation of canopy transpiration rate by applying a plant hormone “abscisic acid”

A method for evaluation of temporal changes in canopy transpiration rate and stomatal conductance in crop fields by using a plant hormone abscisic acid (ABA) has recently been developed. The method was applied to a corn canopy at different growth stages in the upper Yellow River basin, China. Diurnal changes in the canopy transpiration rate and stomatal conductance were evaluated at the initial stage with a leaf area index (LAI) of 0.37 on June 7 and the crop development stage with an LAI of 4.39 on July 15, 2005. The proportions of the accumulated transpiration rate during daytime to the accumulated evapotranspiration were 24% and 74% at the initial and crop development stages, respectively. Stomatal conductance varied in parallel with transpiration rate in the initial stage of the crop. However, in the crop development stage with low soil water content, stomatal conductance reached the maximum value at 10:00 a.m. and thereafter decreased rapidly at around noon with high evaporative demand to corn canopy. This shows the midday stomatal closure was caused by excessive water stress to corn canopy in the crop development stage. Thus, the proposed method with ABA application is useful for evaluation of temporal changes in transpiration rate and stomatal conductance, and hence, can detect the plant water stress.     

Direct and indirect effects of elevated CO2 on transpiration from Quercus myrtifolia in a scrub-oak ecosystem

Elevated atmospheric carbon dioxide (C_a) usually reduces stomatal conductance, but the effects on plant transpiration in the field are not well understood. Using constant‐power sap flow gauges, we measured transpiration from Quercus myrtifolia Willd., the dominant species of the Florida scrub‐oak ecosystem, which had been exposed in situ to elevated C_a (350 µmol mol ^{? 1} above ambient) in open‐top chambers since May 1996. Elevated C_a reduced average transpiration per unit leaf area by 37%, 48% and 49% in March, May and October 2000, respectively. Temporarily reversing the C_a treatments showed that at least part of the reduction in transpiration was an immediate, reversible response to elevated C_a. However, there was also an apparent indirect effect of C_a on transpiration: when transpiration in all plants was measured under common C_a, transpiration in elevated C_a‐grown plants was lower than that in plants grown in normal ambient C_a. Results from measurements of stomatal conductance (g_s), leaf area index (LAI), canopy light interception and correlation between light and g_s indicated that the direct, reversible C_a effect on transpiration was due to changes in g_s caused by C_a, and the indirect effect was caused mainly by greater self‐shading resulting from enhanced LAI, not from stomatal acclimation. By reducing light penetration through the canopy, the enhanced self‐shading at elevated C_a decreased stomatal conductance and transpiration of leaves at the middle and bottom of canopy. This self‐shading mechanism is likely to be important in ecosystems where LAI increases in response to elevated C_a.     

Characteristics of canopy evapotranspiration from a small heterogeneous grassland using thermal imaging

We used thermal imaging in conjunction with the eddy covariance technique to characterize canopy evapotranspiration (ET) from a small heterogeneous grassland. We compared ET estimated by a simple soil–vegetation–atmosphere transfer (SVAT) at field scale (a few 100 m²) with that estimated by the eddy covariance method. These two independent estimates of ET showed a good correlation when the flux source area was the same. However, whereas the eddy covariance method yielded integrated results over a large, variable landscape area, the SVAT model primarily yielded values reflecting just the grassland area. We estimated mapped transpiration (Tr) at a point scale (1 m²) and showed that Tr increased linearly with increasing leaf area index (LAI). Although stomatal conductance of C₃ plants was appreciably larger than that of C₄ plants at the leaf scale, this difference was not reflected in Tr at the canopy scale. Tr may be more sensitive to aerodynamic conditions (wind speed and radiation) or environmental heterogeneity (soil–water) than stomatal conductance. The SVAT model clarified variations in the spatial distribution of Tr over a heterogeneous grassland.     

Coordination of stomatal,hydraulic, and canopy boundary layer properties: Do stomata balance conductances by measuring transpiration?

A striking coordination is observed in sugarcane between prevailing levels of stomatal opening and the hydraulic capacity of the soil, roots and stem to supply the leaves with water. This coordination of vapor phase and liquid phase conductances is associated with decreases in stomatal conductance on a leaf area basis that compensate for increasing leaf area during canopy development, causing transpiration to approach a maximum value on a per plant or ground area basis rather than increase linearly with leaf area. The resulting balance between water loss and water transport capacity maintains leaf water status remarkably constant over a wide range of plant. sizes and growing conditions. These changes in stomatal conductance during development are determined by changes in the composition of the xylem sap rather than by changes in leaf properties. Changes in boundary layer conductance resulting from non-developmental changes in canopy structure such as loding cause additional changes in stomatal conductance mediated by altered humidity at the leaf surface. These maintain a constant level of total canopy vapor phase conductance (stomatal and boundary layer in series) and a constant level of canopy transpiration. These patterns indicate that stomata exert an active role in regulating transpiration even in dense canopies. This control function is consistent with stomatal metering of transpiration, mediated by fluxes of root-derived materials in the xylem sap.     

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。     

Effects of Boundary Layer Conductance on Leaf Gas Exchange

Using an improved gas-exchange technique for leaf chamber the authors' conclusions derived from electrical analogy analysis and simulation have been tested. In most devices for gas-exchange measurements, a fixed ventilation speed is used, which results in a fixed boundary layer conductance of leaf, but the results of experiments are often used to predict canopy transpiration or photosynthesis where the boundary layer conductance changes with the position of the leaf in the canopy and the wind speed above the canopy. To change the boundary layer conductance of a leaf, a barrier of variable size was inserted into the leaf chamber to decrease the wind speed over the leaf. The responses of stomatal conductance, net photosynthetic rate, and transpiration rate to light were then measured. The relationships amongst them have been tested. The experimental results matched well with the results predicted by electrical analogy analysis and simulation in most cases.     

Leaf Response to Water Deficits in Soybeans

Soybeans [Glycine max (L.) Merrill cv. Wayne] plants were subjected to an extended drying cycle in the field to investigate the leaf sensitivity to water deficits. Soybeans in irrigated plots were superior to those in non-irrigated plots in the average size and number of leaflets per plant. Apparent differences in the leaf area distributions in the canopy seemed to be mediated by moisture stress effects associated with leaf senescence and light penetration in the lower depths of the canopy. A major decrease in leaf enlargement occurred near a leaf-water potential of -8 bars, and at - 12 bars, the growth was completely halted. Similar decreases were observed at a stomatal conductance of 0.4 cm/s and at 0.2 cm/s no enlargement was observed.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

华北山区侧柏冠层气孔导度特征及其对环境因子的响应

冠层气孔导度(g_s)是衡量冠层-大气界面水汽通量的重要生物学常数,研究其特征及对环境因子的响应,能为开展森林冠层水汽交换过程的机理性研究提供理论依据.于2014年利用SF-L热扩散式探针测定了侧柏的树干液流密度(J_s),同步监测光合有效辐射(PAR)、饱和水汽压差(VPD)、气温(T)等环境因子,计算侧柏的冠层气孔导度特征并分析其对各环境因子的响应.结果表明: 侧柏液流密度的日变化总体呈双峰曲线,生长季高于非生长季,且胸径越大液流密度越大;冠层气孔导度日变化与单位叶面积冠层蒸腾(E_L)趋势相近,均呈双峰曲线,生长季的冠层气孔导度和蒸腾较非生长季略高.侧柏冠层气孔导度与空气温度呈抛物线关系,在10 ℃左右冠层气孔导度达到峰谷;光合有效辐射以400 μmol·m^-2·s^-1为界,小于该阈值两者呈正相关关系,大于该阈值则冠层气孔导度受其影响较小;与饱和水汽压差呈负对数函数关系,随饱和水汽压差增大而逐渐降低.较高的空气温度和光合有效辐射、较低的饱和水汽压差有利于侧柏形成较大的冠层气孔导度,进而促进冠层蒸腾.     

Study of conductance changes of bilayer lipid membrane induced by electric field

Changes in the bilayer lipid membrane (BLM) conductance induced by electric field were studied. BLMs were formed from diphytanoylphosphocholine (DPhPC) solution in squalene. Certain time after a constant voltage (200-500 mV) was applied to the BLM in the voltage-clamp mode, the BLM conductance started to grow up to approximately 10 nS until the BLM ruptured. The conductance often changed abruptly (with the front duration of less than 33 micros) and then stabilized for a relatively long time (up to 10; 300 ms on average) thus resembling the ion channel activity. The mean amplitude of conductance steps was 650 pS. However, in some cases a slow conductance drift was recorded. When N-methyl-D-glucamine/glutamate ions were used instead of KCl, the conductance changes became 5 times smaller. We suggest that formation in the BLM of single pores approximately 1 nm in diameter should result in the observed changes in BLM conductance. The BLM conductance growth was due to consecutive opening of several such pores. When the electric field amplitude was abruptly decreased (down to 50-100 mV), the conductance dropped rapidly to the background value. When we increased the voltage again, the BLM conductance right after the increase depended on the time BLM spent under "weak" electric field. If this time exceeded 500 ms, the conductance was at the background level, but when the time was diminished, the conductance reached the value recorded before the voltage decrease. These data imply that the closure of the pores should lead to the formation in BLM of small defects (prepores) that can be easily transformed into pores when the voltage is increased. The lifetimes of such prepores did not exceed 500 ms.