根源信号参与调控气孔行为的机制及其农业节水意义

在土壤干旱情况下,根源信号一方面向植物地上部分的长距离传输,为地上部分提供了土壤水分获取能力的测度,另一方面调控气孔开度,抑制蒸腾作用并提高植物的水分利用效率．文中综述了根源信号参与调控植物水分利用的生理机制和理论模型,指出该模型与根系吸水模型、气孔导度模型耦合,能够更好地反映植物叶片对土壤干旱以及大气干旱的响应、评述了在根源信号参与调控植物水分关系的基础上发展的调亏灌溉(RDI)、部分根系干旱(PRD)和控制性交替灌溉(CAI)等有效灌溉手段,有助于合理配置根系层供水量,通过根土相互作用和信号物质的传输,降低蒸腾和提高水分利用效率、另外,根源信号在调控根系生长发育、延缓地上部分生长以调节根冠比例,优化资源分配以利于生殖生长等方面均有所为,为全面提高农田水分利用效率提供节水生理基础。     

Reconciling the optimal and empirical approaches to modelling stomatal conductance

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

Genetic variability of photosynthesis and water use in Balearic grapevine cultivars

A comparison of photosynthetic characteristics of 20 cultivars of grapevine ( Vitis vinifera L. ) from Mallorca (Balearic Islands, Spain) and two widespread cultivars, Cabernet Sauvignon and Chardonnay, was made under irrigation as well as in response to drought. Although these cultivars share a common origin, a high variability was found for several photosynthetic characters under irrigation. Interestingly, these variations were significant for gas-exchange parameters (net CO₂ assimilation, stomatal conductance and intrinsic water use efficiency) but not for chlorophyll fluorescence parameters (maximum photochemical efficiency, electron transport rate and non-photochemical quenching). Since water stress is the most limiting factor for plant production under the Mediterranean climate, it is presumable that these findings reflect specific selection pressures over physiological characteristics related to a balance between net carbon gain and water use. Some cultivars presented high carbon assimilation at the expense of a high water loss, whereas others were water savers, accompanied by low CO₂ assimilation even under irrigation. Escursach was found to be an interesting cultivar, presenting low water consumption at the same time as reasonably high carbon assimilation. These cultivars also showed different responses to drought, which allowed their classification in two main groups: alarmist cultivars, which showed strong reductions of stomatal conductance in response to relatively low decreases of leaf water potential, and luxurious cultivars, showing low reductions of stomatal conductance under water stress.     

Physiological and productive responses of Olea europaea L. cultivars Frantoio and Leccino to a regulated deficit irrigation regime

Abstract

Olive is a drought-tolerant species and it is known that it responds efficiently to any additional water up to a limit. A field experiment was planned with the following aims: to provide estimates of crop evapotranspiration (ETc) to improve water use efficiency during the growing season; to present guidelines for efficient management of irrigation scheduling; and to characterize the relationship between plant water status and optimum fruit yield. These relationships were monitored during four years by analysing the influence of deficit irrigation strategies on mature modern-trained olive trees of cultivars Frantoio and Leccino. Treatments were a non-irrigated control (rain-fed) and three treatments that received a seasonal water amount equivalent to 33, 66 and 100% of ETc, from the beginning of pit hardening to early fruit veraison. Results of the relationship between leaf water potential and maximum stomatal conductance (Ψ_pd vs. g _smax) showed that the stomatal apparatus in Frantoio was more sensitive to water deficit than that of Leccino. Differences in yield between treatments were mainly related to mean fruit weight, indicating that water availability might have affected growing conditions before flowering or during the early stages of fruit growth rather than later in the summer season. Vegetative development was a function of water available to plants. Frantoio achieved the highest crop production per unit of water consumption. Oil quality was scarcely affected by deficit irrigation. Regulated deficit irrigation of olive trees after pit hardening could be recommended, at least under the experimental conditions of this study. Given the different long-term watering response of Frantoio and Leccino, a cultivar-specific irrigation scheduling is advisable.     

Regulated deficit irrigation in different phenological stages of potted geranium plants: water consumption, water relations and ornamental quality

The irrigation water requirements and sensitivity to water deficits of ornamental plants is of great interest to horticultural producers for planning irrigation strategies. The effect of different deficit irrigation strategies on physiological and morphological parameters in geranium plants was studied in different growth phases to evaluate how such strategies can be safely used and to ascertain whether the flowering phase is sensitive to deficit irrigation. Pelargonium × hortorum L.H. Bailey plants, grown in a controlled growth chamber, were subjected to four irrigation treatments: control (100 % water field capacity throughout the experiment), sustainable deficit irrigation (75 % water field capacity throughout the experiment), and two regulated deficit irrigation treatments that included water stress during the vegetative growth phase or during the flowering development phase. Although the total amount of irrigation water was similar in the three deficit irrigation treatments (around 80 % of the control value), the lowest values for both height and flowering were found when deficit irrigation was applied during flowering. This indicates that plant quality does not only depend on the amount of water applied but also on the time when the reduction is applied, and that flowering is the most sensitive phase to water stress. Evapotranspiration was related to the formation of inflorescences and to increased plant height. When the irrigation strategy was changed, plants increased or decreased their water consumption and stomatal conductance to adjust to the new conditions by regulating stomatal opening, although, in general, the values of both parameters remained below those observed in the control plants.     

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。     

夏玉米苗期有限水分胁迫拔节期复水的补偿效应

以作物调亏灌溉原理为基础 ,对夏玉米苗期水分胁迫拔节期复水进行了试验研究。结果表明 ,复水增加了夏玉米叶片气孔导度和光合速率 ,提高叶片水平上的水分利用效率(WUE)。复水 2天后 ,叶片气孔导度和光合速率即恢复到对照水平 ,部分时段 ,特别在下午 ,复水处理表现出高于对照的“反冲”现象。复水使苗期受旱的夏玉米株高、叶面积、地上 (下 )部分干物重和根系生长发育都恢复到或接近充分供水的植株生长水平。使其产量及构成因子与对照接近 ,水分利用效率显著地提高了 2 4 7%。这为玉米中后期的水分管理 ,提高玉米水分利用效率 ,提供了一定的理论依据 ;也提供了一种便于广大农民掌握的简单易行的灌溉方式     

Modelling the effect of fruit growth on surface conductance to water vapour diffusion

BACKGROUND AND AIMS: A model of fruit surface conductance to water vapour diffusion driven by fruit growth is proposed. It computes the total fruit conductance by integrating each of its components: stomata, cuticle and cracks. METHODS: The stomatal conductance is computed from the stomatal density per fruit and the specific stomatal conductance. The cuticular component is equal to the proportion of cuticle per fruit multiplied by its specific conductance. Cracks are assumed to be generated when pulp expansion rate exceeds cuticle expansion rate. A constant percentage of cracks is assumed to heal each day. The proportion of cracks to total fruit surface area multiplied by the specific crack conductance accounts for the crack component. The model was applied to peach fruit (Prunus persica) and its parameters were estimated from field experiments with various crop load and irrigation regimes. KEY RESULTS: The predictions were in good agreement with the experimental measurements and for the different conditions (irrigation and crop load). Total fruit surface conductance decreased during early growth as stomatal density, and hence the contribution of the stomatal conductance, decreased from 80 to 20 % with fruit expansion. Cracks were generated for fruits exhibiting high growth rates during late growth and the crack component could account for up to 60 % of the total conductance during the rapid fruit growth. The cuticular contribution was slightly variable (around 20 %). Sensitivity analysis revealed that simulated conductance was highly affected by stomatal parameters during the early period of growth and by both crack and stomatal parameters during the late period. Large fruit growth rate leads to earlier and greater increase of conductance due to higher crack occurrence. Conversely, low fruit growth rate accounts for a delayed and lower increase of conductance. CONCLUSIONS: By predicting crack occurrence during fruit growth, this model could be helpful in managing cropping practices for integrated plant protection.     

宁夏平原北部地下水埋深浅地区不同灌木的水分来源

宁夏平原北部引黄灌区地下水埋深浅是该地区土壤盐碱化的主要原因, 种植耐盐植物可以吸收利用地下水, 在降低地下水位的同时可以减少对地面灌溉的依赖。为了分析银川平原北部4种灌木对不同水源的利用特征, 于2010年生长季测定了灌溉前后20年生多枝柽柳(Tamarix ramosissima)、3年生多枝柽柳、3年生宁夏枸杞(Lycium barbarum)和3年生四翅滨藜(Atriplex canescens)木质部水及不同潜在水源稳定氧、氢同位素组成(δ¹⁸O和δD), 应用IsoSource同位素线性混合模型估算了不同灌木对不同水源的利用率。同时测定了0-200 cm土壤剖面的全盐含量、含水量和pH值以及灌溉前后光合气体交换参数。结果表明: 不同深度土壤水δ¹⁸O和δD值存在较大差异, 并呈规律性变化。土壤水δ¹⁸O和δD值随深度加深呈逐渐降低的趋势。灌溉后80 cm以上土壤水δ¹⁸O和δD值低于灌溉前。无论灌溉前还是灌溉后, 20年生多枝柽柳与3年生灌木相比具有更低的δ¹⁸O和δD值。灌溉前, 3年生多枝柽柳、宁夏枸杞和四翅滨藜主要利用表层土壤水(70.1%、52.3%和48.9%); 20年生多枝柽柳对地下水的利用率最高(21.5%)。灌溉后, 3年生多枝柽柳和宁夏枸杞对80-140 cm土壤水利用率较高(59.5%和58.8%)。20年生多枝柽柳对地下水的利用率最高(18.3%)。灌溉前, 20年生多枝柽柳净光合速率、气孔导度和蒸腾速率显著高于其他3种灌木, 灌溉后3年生四翅滨藜净光合速率最高。灌溉对3年生多枝柽柳和宁夏枸杞的净光合速率和气孔导度有显著影响。无论灌溉前还是灌溉后, 3年生四翅滨藜瞬间水分利用效率均高于其他3种灌木。研究表明, 不同灌木在不同水分条件下水分利用策略不同, 这主要与植物种类及树龄有关。灌溉前幼龄多枝柽柳凭借其对干旱较强的忍耐能力利用浅层不饱和土壤水, 灌溉后其又转而利用中层土壤水, 表现出潜水湿生植物的特征, 主要吸收利用深层土壤水分, 对灌溉反应不明显。     

Response of Cynodon dactylon to prolonged water deficits under saline conditions

Competition for available water resources in both arid and semi-arid locations has led to greater scrutiny of turfgrass irrigation. Irrigation management strategies, including deficit irrigation, need further investigation. The objective of this research was to determine the physiological response of bermudagrass (Cynodon dactylon (L.) Pers.) to prolonged water deficits under saline conditions. Bermudagrass was grown in large columns packed with three different soil types (sandy loam, silt loam and clay). Synthesized saline irrigation water was applied at three different salinity levels (1.5, 3.0 and 6.0 dS/m). Two previous experiments that were conducted with these columns over a 3.5 year period led to differential profile salinization in all 27 columns. At the end of this 3.5 year period, all irrigations were terminated and plant growth and water status were monitored over a 95 day dry-down period. Midday stomatal conductance, leaf water potential, canopy temperature, soil water in storage and stolon elongation were measured over the experimental period. On day 95, above ground tissue was harvested for dry weight and elemental tissue analysis. Midday stomatal conductance decreased around day 30 in all columns regardless of soil salinity. This decrease was not associated with a threshold leaf water potential, as midday leaf water potentials remained constant over a 60 day period. Stolon elongation also ceased before any deviation in the midday leaf water potential occurred. A concomitant reduction in evapo ranspiration was not associated with the measured decrease in stomatal conductance. This would suggest that bermudagrass may have regulated stomatal activity to compensate for lower conductances during periods of greater stress and/or that measured midday stomatal conductances cannot always be directly coupled to extended time evapotranspiration measurements.     

Plant water-regulation strategies: Isohydric versus anisohydric behavior

Water is a vital resource for plant survival, growth and distribution, and it is of significance to explore mechanisms of plant water-relations regulation and responses to drought in ecophysiology and global change ecology. Plants adapt to different climates and soil water regimes and develop divergent water-regulation strategies involving a suite of related traits, of which two typical types are isohydric and anisohydric behaviors. It is critical to distinguish water-regulation strategies of plants and reveal the underlying mechanisms for plant breeding and vegetation restoration especially in xeric regions; and it is also important for developing more accurate vegetation dynamic models and predicting vegetation distribution under climate change scenarios. In this review, we first recalled the definitions of isohydric and anisohydric regulations and three quantitative classification methods that were established based on the relationships (1) between stomatal conductance and leaf water potential, (2) between stomatal conductance and vapor pressure deficit, (3) between predawn and midday leaf water potentials. We then compared the two water-regulation strategies in terms of hydraulics and carbon-economics traits. We synthesized the mechanisms of plant water-regulation and found that the interaction between hydraulic and chemical signals was the dominant factor controlling plant water-regulation behavior. Last, we proposed three promising aspects in this field: (1) to explore reliable and universal methods for classifying plant water-regulation strategies based on extensive investigation of the traits related with plant water-relations in various regions; (2) to explore relationships between plant water-regulation strategies and traits of hydraulics, morphology, structure, and function in order to provide reliable parameters for improving vegetation dynamic models; and (3) to deeply understand the processes of plant water-regulation at different spatial and temporal scales, and reveal mechanisms of plants’ responses and adaption to environmental stresses (especially drought).     

Stomatal numbers of soybean and response to water stress

The relationship among stomatal density, photosynthetic rate, leaf conductance, plant growth, bean yield and kaempferol triglucoside (K9) in the leaves of soybean (Glycine max (L.) Merr.) was examined in two field tests. K9 in the leaves was associated with reduced stomatal density, reduced photosynthetic rate, reduced stomatal conductance, reduced plant weight and lower bean yield. Plants with high stomatal frequency (lacking K9) were better able to take advantage of increased water supply by increasing stomatal conductance (upper surface), transpiration and bean yield. Plants with low stomatal frequency (with K9) were unresponsive to irrigation and in this sense were more tolerant of water stress, but their overall yield was low.     

Water relations of yellow sweetclover under the synergy of drought and selenium addition

In the last two decades drought and elevated toxic metal concentration phenomena in plants have gained the interest of the scientific world. Nevertheless, up to day relatively little ecophysiological research concerning the effect of water stress and elevated selenium (Se) concentration on plant water relations is available. A pot experiment was conducted in order to evaluate the effects of the implied synergy of drought and Se uptake on water relations of yellow sweetclover (Melilotus officinalis L.). The effects of two different Se concentrations (0 mg Se L^?1 irrigation water, 3 mg Se L^?1 irrigation water) and two water regimes (full irrigation — limited irrigation) applied to seedlings of yellow sweetclover were detected by measuring changes in water potential, relative water content, stomatal conductance, transpiration rate and tissue Se concentration. The findings suggest that yellow sweetclover, concentrating up to 200 μg Se g^?1 dry weight in its tissues, could be considered a secondary Se accumulator. Se effect on water relations was more evident under limited irrigation, as expressed by decreased values of leaf water potential, transpiration rate and stomatal conductance, limiting the flux rate of the water solution in the conducting system of the plant.     

The efficiency of water use in water stressed plants is increased due to ABA induced stomatal closure

Gas exchange and abscisic acid content of Digitalis lanata EHRH. have been examined at different levels of plant water stress. Net photosynthesis, transpiration and conductance of attached leaves declined rapidly at first, then more slowly following the withholding of irrigation. The intercellular partial pressure of CO₂ decreased slightly. The concentration of 2-cis(S)ABA increased about eight-fold in the leaves of non-irrigated plants as compared with well-watered controls. A close linear correlation was found between the ABA content of the leaves and their conductance on a leaf area basis. In contrast, the plot of net assimilation versus ABA concentration was curvilinear, leading to an increased efficiency of water use during stress. After rewatering, photosynthesis reached control values earlier than transpiration, leaf conductance and ABA content. From these data it is concluded that transpiration through the stomata is directly controlled by the ABA content, whereas net photosynthesis is influenced additionally by other factors.Possible reasons for the responses of photosynthesis and water use efficiency to different stress and ABA levels are discussed.Abbreviations A net CO₂ assimilation - ABA abscisic acid - C_i intercellular CO₂ concentration - g stomatal conductance - T transpiration - WUE water use efficiency     

叶肉细胞导度研究进展

叶肉细胞导度指叶片叶肉细胞内部的CO2扩散能力,在植物生理生态及全球气候变化和陆地生态系统相互关系的研究中具有重要作用。系统介绍了叶肉细胞导度的发现、发展过程及其研究进展、几种目前国际上常用的叶肉细胞导度测度方法的原理、计算过程;强调了叶肉细胞导度作为光合作用扩散过程一部分的重要意义,明确了叶肉细胞导度的定义及分布范围。并探讨了不同方法的优缺点及注意事项。总结分析了叶肉细胞导度对不同环境因子(温度、水分及环境中CO2和O3浓度等)的响应,从不同角度对叶肉细胞导度的生态学意义进行了简单的概括。对叶肉细胞导度的未来研究进行了展望。     

植物气孔气态失水与SPAC系统液态供水的相互调节作用研究进展

讨论了植物气孔气态失水与SPAC系统液态供水相互作用研究领域的一些重要现象和行为.当植物水力信号和化学信号共同作用促进气孔对叶水势的调节时,植物对叶水势的调节表现为等水行为.气孔对环境湿度变化响应的反馈机制可用来解释土壤干旱条件下气孔和光合的午休现象,以及气孔导度和水流导度之间的相关关系;而气孔对环境湿度变化响应的前馈机制,则可用来解释气孔导度对大气 叶片间水汽饱和差的滞后反应.植物最大限度地利用木质部传输水分的策略,要求气孔快速响应以避免木质部过度气穴化和短时间内将气穴逆转的相应机制.     

桃树冠层蒸腾动态的数学模拟

将气孔导度公式、Penman—Monteith公式和土壤水分限制模型相结合,可以模拟出不同环境因子对植物蒸腾进程的影响。通过对盆栽桃树（Prunus persica var．nectadna Maxim．）数值模拟发现：影响桃树蒸腾速率的主要气象因子是太阳辐射、大气温度和湿度。植物通过气孔导度的改变来响应气象因子的变化,蒸腾的日变化主要是由气象因子的日变化引起的。土壤的水分状况也对气孔导度有显著的影响,进而影响植物的蒸腾大小。通过数值模拟还发现植物的蒸腾量并不总是随叶面积的增大而增大,对于桃树而言叶面积指数为4左右时日蒸腾量达到最大值。通过对气孔导度和蒸腾速率的模拟值和实测值进行检验发现,两者基本吻合,说明利用数学模拟的方法可以求出不同环境条件和不同叶面积桃树冠层的蒸腾速率。     

Exploring thermal imaging variables for the detection of stress responses in grapevine under different irrigation regimes

Temperatures of leaves or canopies can be used as indicators of stomatal closure in response to soil water deficit. In 2 years of field experiments with grapevines (Vitis vinifera L., cvs Castel?o and Aragonês), it was found that thermal imaging can distinguish between irrigated and non-irrigated canopies, and even between deficit irrigation treatments. Average canopy temperature was inversely correlated with stomatal conductance measured with a porometer. Variation of the distribution of temperatures within canopies was not found to be a reliable indicator of stress. A large degree of variation between images was found in reference 'wet' and 'dry' leaves used in the first year for the calculation of an index proportional to stomatal conductance. In the second year, fully irrigated (FI) (100% Et(c)) and non-irrigated (NI) canopies were used as alternatives to wet and dry leaves. A crop water stress index utilizing these FI and NI 'references', where stressed canopies have the highest values and non-stressed canopies have the lowest values, was found to be a suitable measure for detecting stress. It is suggested that the average temperatures of areas of canopies containing several leaves may be more useful for distinguishing between irrigation treatments than the temperatures of individual leaves. Average temperatures over several leaves per canopy may be expected to reduce the impact of variation in leaf angles. The results are discussed in relation to the application of thermal imaging to irrigation scheduling and monitoring crop performance.     

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.     

黄土高原地区春小麦对有限灌溉的反应及其生理生态基础

从对黄土高原地区有限灌溉条件下作物生理生态反应的众多研究中得出：（1）水分轻度亏缺时,作物可通过根信号物质ABA调节叶片的气孔导度。非水力根信号作用太强,可因降低光合作用而减少干物质生产和影响干物质分配模式而影响产量和水分利用效率,故削弱非水力根信号的作用将有利于提高产量。（2）浅层根系占根系总量比值越高,对干旱越敏感,表现为根信号能力增强;深层根系所占比例越高,越有利于土壤深层水分利用,并可削弱根信号,同理,给土壤中下层补水或采用播种前灌溉,可因为减少了无效蒸发,且削弱根信号而提高水分利用率。（3）本地区有限灌溉的最佳时期由于降水变率较高而变得较为复杂,不同降水年型,最佳灌溉时期差异很大,对有限灌溉进行科学管理还需要做更多的研究工作。