水分再分配对土壤-植物系统养分循环的生态意义

水分再分配(hydraulic redistribution, HR)作为一个普遍存在的生物物理过程, 在缓解植物干旱胁迫、调节植物种间关系和群落组成、影响生态系统水碳平衡等方面具有重要的生态意义。近年来, 同位素标记示踪技术的应用促进了HR的深入研究, 该文综述了HR对土壤-植被系统养分循环的影响。HR能改善干燥土层的水分状况, 防止根系栓塞, 促进细根存活与生长, 提高微生物活性, 从而促进植物对表层土壤养分(尤其是氮)的吸收; HR还通过水分下传作用促进植物对深层土壤中磷和金属离子的吸收。HR促进土壤养分库的上下交换与流动, 调节植物与土壤的氮磷比, 因此其影响可能具有全球意义。在全球变化(如氮沉降)背景下, 有必要深入探索HR在生物地球化学循环过程中的影响和作用, 并将其纳入生态系统模型中。     

植物-土壤系统中水分再分配作用研究进展

在过去100多年里,植物与土壤之间水分关系的研究多侧重于植物本身的水分利用方式、水分利用效率及其植物根系对水分的吸收等.然而进入20世纪80年代后期,植物生理生态学研究人员开始将注意力转移到植物根系对土壤水分的调节作用,即水分再分配（hydraulic redistribution）作用,具体地讲是在水势差的驱动下水分由根系向土壤中释出的一种双向和被动的水分运转过程,其中既包含水分由深层土壤向表层土壤的释出,也包括由表层土壤向深层土壤的流动,同时还涵盖了水分在水平方向上的侧向运输过程.伴随着研究手段的不断提高和对生态系统水平衡问题的关注,水分再分配逐渐成为近代植物生态学和水文学的交叉学科生态水文学（ecohydrology）的核心研究内容之一.目前该领域的研究已经阐明水分再分配作用在不同程度上对植物个体蒸腾、碳同化速率有很大贡献,有利于提高根系生活力和土壤养分;另外,在不断扩展的生态系统生态学研究中,也加强了对制约水分再分配作用发生的外部因子的认识.回顾和分析了水分再分配的研究历史、生态学意义、影响因素、测定方法等,特别提出阐述浅根系植物对水分再分配作用的依赖性与依赖程度,从植物进化学角度解释水分再分配作用发生的生理学基础和意义及水分再分配作用对土壤微生物活性的影响等方面将是未来研究的几个重点方向.     

共生条件下三种荒漠灌木的根系分布特征及其对降水的响应

以全根系挖掘法,对共生于原始盐生荒漠生境中的多枝柽柳[Tamarix ramosissima (Ledeb.)]、梭梭[Haloxylon ammodendron(C. A. Mey.)Bunge]、琵琶柴[Reaumuria soongorica (Pall.) Maxim.]的根系分布特征进行了研究;对降水引发的湿润-干旱周期中植物同化枝水势、蒸腾速率的变化过程进行了跟踪观测,并据此计算3种植物的水分胁迫效应指数和土壤-植物系统导水度,以最终确定3种植物用水策略和其对降水的响应特征.研究结果表明,多枝柽柳的吸收根系分布范围从地下50cm到310cm,单株平均总吸收根表面积为30249.2cm2;梭梭的根系分布范围0～250cm,单株平均总吸收根表面积12847.3 cm2;琵琶柴的根系分布范围0～80cm,单株平均总吸收根表面积361.8 cm2.多枝柽柳为深根植物,主要利用地下水和深层土壤水,在降水引发的湿润-干旱周期中,其植物水分生理参数对降水无响应.琵琶柴为浅根植物,对降水响应极为显著.梭梭的根系分布特征介于多枝柽柳和琵琶柴之间,对地下水和降水都有利用,对降水响应显著.3种荒漠灌木对降水的响应差异显然与其根系分布、水分利用策略密切相关,在未来降水发生变化的情景下,根系分布特征的差异将决定着植物在水分资源竞争中的地位.具有较强根系形态可塑性的物种,如梭梭,将具有明显的竞争优势.     

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。     

土壤水分与温度共同作用对植物根系水分传导的效应

 本文根据不同大气环境温度和土壤温度及不同土壤含水率处理条件下的玉米、向日葵、台湾相思(Acacia confusa)、银合欢(Leucaena glauca)的试验资料，分析了土壤水分和温度以及土壤水分与温度共同作用对植物根系水分传导的效应。台湾相思和银合欢的试验结果表明，在一定的土壤水分范围内，高温(白天/夜晚的温度为40／30℃)环境中的根系水分传导大于低温(30/25℃)环境中的，但当根系水分胁迫十分严重(台湾相思根系水势小于-1.5MPa，银合欢根系水势小于-2.0MPa)时，30/25℃环境的根系水分传导反而大于40／30℃环境的；玉米、向日葵的试验结果表明，在一定土壤温度范围内，根系水分传导随土壤温度增加而增加，其增加的幅度与生育阶段有关；在向日葵生育期土壤温度高于35℃、玉米生育期高于30℃时，其根系水分传导随温度增加而降低。通过植物根区土壤逐渐干旱和干旱复水后的试验，其结果表明复水后根系水分传导上升较快，银合欢复水1.5d、向日葵复水3d后测得的根系水分传导即可达到受旱前的水平，其后的水分传导还略高于一直充分供水处理的，表明根系经受一定程度的干旱锻炼后，对其水分传导具有明显的补偿效应。在干旱和复水过程中根系水分传导与根水势的变化规律相一致。     

水分对土壤呼吸的影响及机理

土壤呼吸是陆地碳循环的重要环节,在全球变化的背景下,研究水分对土壤呼吸的影响,能为探索陆地生态系统在碳循环方面的源—汇功能和揭示碳的失汇之迷提供有力的证据。综述了水分对土壤呼吸的影响及其机理。土壤呼吸是一个复杂的生态学过程,大气降水对土壤呼吸的影响结果是因时、因地而异,在湿润的生态系统或者干湿交替的生态系统中比较湿润的季节．降水事件对土壤呼吸可能会产生比较明显的抑制现象;而在干旱的生态系统或有干湿交替季节的生态系统中比较干旱的季节里,降水事件可能会强烈地激发土壤呼吸。其对土壤呼吸的影响机理包括水分对土壤孔隙中CO2替代、对CO2扩散的阻滞、对微生物活动的刺激和对微生物生物量的影响等。由于实验方法和标准的不一致以及影响土壤呼吸的因素的多样性。水分量的变化对土壤呼吸的影响很难以一个统一的方程来描述,总的来说,最优的水分状况通常是接近最大田间持水力,当土壤处于过于或过湿状态时,土壤呼吸会受到抑制。水分量的变化对土壤呼吸的影响机制在于可溶性有机质、土壤的通透性、微生物与植物根系生命活动等都随土壤水分状况不同而发生相应的改变。关于水分与土壤呼吸的关系研究今后应该主要集中在：(1)水分对根系呼吸和土壤微生物呼吸分别产生的影响;(2)全球变化后水分格局的变化对全球陆地生态系统土壤呼吸格局的潜在影响;(3)人类活动通过直接或间接改变水分状况而对土壤释放CO2的贡献率。     

土壤深层供水对冬小麦干物质生产的影响

采用根系研究装置研究了土壤深层供水对冬小麦干物质生产的影响 .结果表明 ,上层低湿度下层高湿度的处理在小麦灌浆期仍然保持了较高的土壤和叶片含水量 ,具有发达的根系 ,特别是 1m以下的根量在 4个处理中为最高 ,旗叶和穗的干重也最大 ,具有最大的产量潜力 .本研究表明 ,上层土壤较干下层土壤湿润有利于发挥小麦根信号的积极作用 ,平衡水分利用 ,同时通过对土壤水分的合理调节可以促进深层根的发育 ,有利于提高产量和水分利用效率 .     

稳定同位素技术在植物水分利用研究中的应用

近20a稳定同位素技术在植物生态学研究中的应用得到了长足发展,使得对植物与水分关系也有了更深一步的了解。介绍稳定同位素性碳、氢、氧同位素在研究植物水分关系中的应用及进展,以期能为国内植物水分利用研究提供参考。由于植物根系从土壤中吸收水分时并不发生同位素分馏,对木质部水分同位素分析有助于对植物利用水分来源,生态系统中植物对水分的竞争和利用策略的研究,更好地了解生态系统结构与功能。稳定碳同位素作为植物水分利用效率的一个间接指标,在不同水分梯度环境中,及植物不同代谢产物与水分关系中有着广泛的应用。同位素在土壤-植被-大气连续体水分中的应用,有助于了解生态系统的水分平衡。随着稳定同位素方法的使用,植物与水分关系的研究将取得更大的进展。     

典型林区水分氢氧稳定同位素在土壤-植物-大气连续体中的分布特征

土壤-植物-大气连续体(SPAC)中水循环是水文学和生态学研究的重要内容,氢氧稳定同位素在不同水体中组成特征的差异可以指示水分循环过程。本研究通过分析成都平原区亚热带常绿阔叶林中降水、土壤水、植物水的同位素组成,探讨SPAC系统中水分的氢氧稳定同位素演化特征,揭示区域水循环不同界面过程。结果表明: 研究区雨季大气降水线方程为: δD=7.13δ¹⁸O+2.35(R²=0.99),土壤蒸发线方程为: δD=6.98δ¹⁸O-0.32(R²=0.92)。在降水→土壤水→植物水的界面水输送过程中,氢氧同位素逐渐富集。浅层土壤(0～35 cm)水δ¹⁸O受降水的直接影响,响应关系明显,中深层土壤(35～100 cm)水则相对稳定。观测期间,植物木质部水同位素比土壤水略微富集,说明水分在植物体内输送过程中可能通过韧皮部或树皮发生轻微蒸发或蒸腾。采用直接相关法初步估计植物对不同土层土壤水的利用情况,樟树主要利用中层土壤水,构树主要利用浅层土壤水,金星蕨因根系分布浅更倾向于利用浅层土壤水和植物截留的降水。与金星蕨相比,樟树和构树的叶片水分蒸发和同位素动力分馏程度更强。     

基于δD和δ18O的青海湖流域芨芨草水分利用来源变化研究

水分条件是限制干旱半干旱地区植物生长重要的生态因子,为了揭示青海湖流域典型生态系统下芨芨草植物的水分利用来源及其如何响应水分条件的变化,选择了自然和干旱控制条件下芨芨草植物,通过测定芨芨草植物茎水和各潜在水源(土壤水、地下水及降水)中δD、δ~(18)O组成,并利用直接比较分析法和多源混合模型计算芨芨草植物对土壤水的利用比例。研究结果表明:表层土壤水分和土壤水中δD、δ~(18)O值表征出较大波动范围,其直接受降水和蒸发作用影响,土壤蒸发线的斜率和截距明显小于大气水线斜率和截距,表明土壤水中同位素组成经历了强烈的蒸发分馏过程,而芨芨草茎水中δD、δ~(18)O值都集中分布土壤水蒸发线附近,说明芨芨草根系主要利用不同深度的土壤水。自然条件下芨芨草在生长季初期(6月)利用表层土壤水(0—10cm,45.1%),8—9月份大降水事件影响土壤含水量和同位素组成,降水入渗深度较深且芨芨草根系对土壤水分吸收的比例相差不大,表明根系在土壤含水量较高时均能吸水不同深度土壤水。在干旱控制条件下芨芨草在7月初主要利用表层土壤水(0-30cm),随着表层土壤水分的减少,根系吸收深度转向较深土壤层,而灌溉后表层土壤水分明显增加,其吸收深度又转向表层,表明芨芨草根系吸收深度能敏感地响应土壤水分的变化。另外还发现芨芨草在生长季内并未直接利用地下水。     

Modelled hydraulic redistribution by sunflower (Helianthus annuus L.) matches observed data only after including night‐time transpiration

The movement of water from moist to dry soil layers through the root systems of plants, referred to as hydraulic redistribution (HR), occurs throughout the world and is thought to influence carbon and water budgets and ecosystem functioning. The realized hydrologic, biogeochemical and ecological consequences of HR depend on the amount of redistributed water, whereas the ability to assess these impacts requires models that correctly capture HR magnitude and timing. Using several soil types and two ecotypes of sunflower (Helianthus annuus L.) in split‐pot experiments, we examined how well the widely used HR modelling formulation developed by Ryel et al. matched experimental determination of HR across a range of water potential driving gradients. H. annuus carries out extensive night‐time transpiration, and although over the last decade it has become more widely recognized that night‐time transpiration occurs in multiple species and many ecosystems, the original Ryel et al. formulation does not include the effect of night‐time transpiration on HR. We developed and added a representation of night‐time transpiration into the formulation, and only then was the model able to capture the dynamics and magnitude of HR we observed as soils dried and night‐time stomatal behaviour changed, both influencing HR.     

The magnitude of hydraulic redistribution by plant roots: a review and synthesis of empirical and modeling studies

Hydraulic redistribution (HR) - the movement of water from moist to dry soil through plant roots - occurs worldwide within a range of different ecosystems and plant species. The proposed ecological and hydrologic impacts of HR include increasing dry-season transpiration and photosynthetic rates, prolonging the life span of fine roots and maintaining root-soil contact in dry soils, and moving rainwater down into deeper soil layers where it does not evaporate. In this review, we compile estimates of the magnitude of HR from ecosystems around the world, using representative empirical and modeling studies from which we could extract amounts of water redistributed by plant root systems. The reported average magnitude of HR varies by nearly two orders of magnitude across ecosystems, from 0.04 to 1.3 mm H(2)O d(-1) in the empirical literature, and from 0.1 to 3.23 mm H(2)O d(-1) in the modeling literature. Using these synthesized data, along with other published studies, we examine this variation in the magnitude of upward and downward HR, considering effects of plant, soil and ecosystem characteristics, as well as effects of methodological details (in both empirical and modeling studies) on estimates of HR. We take both ecological and hydrologic perspectives.     

Hydraulic redistribution affects modeled carbon cycling via soil microbial activity and suppressed fire

Hydraulic redistribution (HR) of water from moist to drier soils, through plant roots, occurs world‐wide in seasonally dry ecosystems. Although the influence of HR on landscape hydrology and plant water use has been amply demonstrated, HR's effects on microbe‐controlled processes sensitive to soil moisture, including carbon and nutrient cycling at ecosystem scales, remain difficult to observe in the field and have not been integrated into a predictive framework. We incorporated a representation of HR into the Community Land Model (CLM4.5) and found the new model improved predictions of water, energy, and system‐scale carbon fluxes observed by eddy covariance at four seasonally dry yet ecologically diverse temperate and tropical AmeriFlux sites. Modeled plant productivity and microbial activities were differentially stimulated by upward HR, resulting at times in increased plant demand outstripping increased nutrient supply. Modeled plant productivity and microbial activities were diminished by downward HR. Overall, inclusion of HR tended to increase modeled annual ecosystem uptake of CO₂ (or reduce annual CO₂ release to the atmosphere). Moreover, engagement of CLM4.5′s ground‐truthed fire module indicated that though HR increased modeled fuel load at all four sites, upward HR also moistened surface soil and hydrated vegetation sufficiently to limit the modeled spread of dry season fire and concomitant very large CO₂ emissions to the atmosphere. Historically, fire has been a dominant ecological force in many seasonally dry ecosystems, and intensification of soil drought and altered precipitation regimes are expected for seasonally dry ecosystems in the future. HR may play an increasingly important role mitigating development of extreme soil water potential gradients and associated limitations on plant and soil microbial activities, and may inhibit the spread of fire in seasonally dry ecosystems.     

Water uptake and hydraulic redistribution across large woody root systems to 20 m depth

Deep water uptake and hydraulic redistribution (HR) are important processes in many forests, savannas and shrublands. We investigated HR in a semi‐arid woodland above a unique cave system in central Texas to understand how deep root systems facilitate HR. Sap flow was measured in 9 trunks, 47 shallow roots and 12 deep roots of Quercus, Bumelia and Prosopis trees over 12 months. HR was extensive and continuous, involving every tree and 83% of roots, with the total daily volume of HR over a 1 month period estimated to be approximately 22% of daily transpiration. During drought, deep roots at 20 m depth redistributed water to shallow roots (hydraulic lift), while after rain, shallow roots at 0–0.5 m depth redistributed water among other shallow roots (lateral HR). The main driver of HR appeared to be patchy, dry soil near the surface, although water may also have been redistributed to mid‐level depths via deeper lateral roots. Deep roots contributed up to five times more water to transpiration and HR than shallow roots during drought but dramatically reduced their contribution after rain. Our results suggest that deep‐rooted plants are important drivers of water cycling in dry ecosystems and that HR can significantly influence landscape hydrology.     

Transverse hydraulic redistribution by a grapevine

Root hydraulic redistribution has been shown to occur in numerous plant species under both field and laboratory conditions. To date, such water redistribution has been demonstrated in two fundamental ways, either lifting water from deep edaphic sources to dry surface soils or redistributing water downward (reverse flow) when inverted soil Ψ_s gradients exist. The importance of hydraulic redistribution is not well documented in agricultural ecosystems under field conditions, and would be important because water availability can be temporally and spatially constrained. Herein we report that a North American grapevine hybrid (Vitis riparia × V. berlandieri cv 420 A) growing in an agricultural ecosystem can redistribute water from a restricted zone of available water under a drip irrigation emitter, laterally across the high resistance pathways of the trunk and into roots and soils on the non-irrigated side. Deuterium-labelled water was used to demonstrate lateral movement across the vine's trunk and reverse flow into roots. Water redistribution from the zone of available water and into roots distant from the source occurred within a relatively short time frame of 36 h, although overnight deposition into rhizosphere soils around the roots was not detected. Deuterium was eventually detected in rhizosphere soils adjacent to roots on the non-irrigated side after 7 d. Application of identical amounts of water with the same deuterium enrichment level (2%) to soils without grapevine roots showed that physical transport of water through the vapour phase could not account for either downward or transverse movement of the label. These results confirmed that root presence facilitated the transport of label into soils distant from the wetted zone. When deuterium-labelled water was allowed to flow directly into the trunk above the root–trunk interface, reverse flow occurred and lateral movement across the trunk and into roots originating around the collar region did not encounter large disproportionate resistances. Rapid redistribution of water into the entire root system may have important implications for woody perennial cultivars growing where water availability is spatially heterogeneous. Under the predominantly dry soil conditions studied in this investigation, water redistributed into roots may extend root longevity and increase the vines water capacitance during periods of high transpiration demand. These benefits would be enhanced by diminished water loss from roots, and could be equally important to other cited benefits of hydraulic redistribution into soils such as enhancement of nutrient acquisition.     

Seasonal variation of water uptake of a Quercus suber tree in Central Portugal

Hydraulic redistribution (HR) is the phenomenon where plant roots transfer water between soil horizons of different water potential. When dry soil is a stronger sink for water loss from the plant than transpiration, water absorbed by roots in wetter soil horizons is transferred toward, and exuded into dry soil via flow reversals through the roots. Reverse flow is a good marker of HR and can serve as a useful tool to study it over the long-term. Seasonal variation of water uptake of a Quercus suber tree was studied from late winter through autumn 2003 at Rio Frio near Lisbon, Portugal. Sap flow was measured in five small shallow roots (diameter of 3–4 cm), 1 to 2 m from the tree trunk and in four azimuths and at different xylem depths at the trunk base, using the heat field deformation method (HFD). The pattern of sap flow differed among lateral roots as soil dried with constant positive flow in three roots and reverse flow in two other roots during the night when transpiration ceased. Rain modified the pattern of flow in these two roots by eliminating reverse flow and substantially increasing water uptake for transpiration during the day. The increase in water uptake in three other roots following rain was not so substantial. In addition, the flux in individual roots was correlated to different degrees with the flux at different radial depths and azimuthal directions in trunk xylem. The flow in outer trunk xylem seemed to be mostly consistent with water movement from surface soil horizons, whereas deep roots seemed to supply water to the whole cross-section of sapwood. When water flow substantially decreased in shallow lateral roots and the outer stem xylem during drought, water flow in the inner sapwood was maintained, presumably due to its direct connection to deep roots. Results also suggest the importance of the sap flow sensor placement, in relation to sinker roots, as to whether lateral roots might be found to exhibit reverse flow during drought. This study is consistent with the dimorphic rooting habit of Quercus suber trees in which deep roots access groundwater to supply superficial roots and the whole tree, when shallow soil layers were dry.     

Water and Nutrient Dynamics in Surface Roots and Soils are not Modified by Short-term Flooding of Phreatophytic Plants in a Hyperarid Desert

Little is known of the mechanisms employed by woody plants to acquire key resources such as water and nutrients in hyperarid environments. For phreatophytic plants, deep roots are necessary to access the water table, but given that most nutrients in many desert ecosystems are stored in the upper soil layers, viable shallow roots may be equally necessary for nutrient uptake. We sought to better understand the interaction between water and nutrient uptake from soil horizons differing in the relative abundance of these resources. To this end, we monitored plant water and nutrient status before and after applying flood irrigation to four phreatophytic perennial plant species in the remote hyperarid Taklamakan desert in western China. Sap flow in the roots of five plants of the perennial desert species Alhagi sparsifolia Shap., Karelina caspica (Pall.) Less., Calligonum caput medusea Schrenk, and Eleagnus angustifolia Hill. was monitored using the heat ratio method (HRM). Additionally we measured predawn and midday water potential, foliar nitrate reductase activity (NRA), xylem sap nutrient concentration and the concentration of total solutes in the leaves before, 12 and 96 h after flooding to investigate possible short-term physiological effects on water and nutrient status. Rates of sap flow measured during the day and at night in the absence of transpiration did not change after flooding. Moderately high rates of sap flow (HRM heat pulse velocity, 5–25 cm h⁻¹) detected during the day in soils that had a near zero water content at the surface indicated that all species had contact to groundwater. There was no evidence from sap flow data that plants had utilised flood water to increase maximum rates of transpiration under similar climatic conditions, and there was no evidence of a process to improve the efficiency of water or nutrient uptake, such as hydraulic redistribution (i.e. the passive movement of water from moist soil to very dry soil via roots). Measurements of plant water status, xylem sap nutrient status, foliar NRA and the concentration of osmotically active substances were also unaffected by flood irrigation. Our results clearly show that groundwater acts as the major source of water and nutrients for these plants. The inability of plants to utilise abundant surface soil–water or newly available nutrients following irrigation was attributed to the absence of fine roots in the topsoil layer.     

Hydraulic redistribution: limitations for plants in saline soils

Hydraulic redistribution (HR), the movement of water from wet to dry patches in the soil via roots, occurs in different ecosystems and plant species. By extension of the principle that HR is driven by gradients in soil water potential, HR has been proposed to occur for plants in saline soils. Despite the inherent spatial patchiness and salinity gradients in these soils, the lack of direct evidence of HR in response to osmotic gradients prompted us to ask the question: are there physical or physiological constraints to HR for plants in saline environments? We propose that build‐up of ions in the root xylem sap and in the leaf apoplast, with the latter resulting in a large predawn disequilibrium of water potential in shoots compared with roots and soil, would both impede HR. We present a conceptual model that illustrates how processes in root systems in heterogeneous salinity with water potential gradients, even if equal to those in non‐saline soils, will experience a dampened magnitude of water potential gradients in the soil–plant continuum, minimizing or preventing HR. Finally, we provide an outlook for understanding the relevance of HR for plants in saline environments by addressing key research questions on plant salinity tolerance.     

干旱区水力提升的生态作用

水力提升是某些植物通过深层根系从较湿的深层土壤中吸取水分再通过浅层根系在较干的浅层土壤中释出的过程。水力提升所释出水量及其释出过程具有积极的生态学意义,它不但为伴生植物提供新的水源、改善其邻近植物的蒸腾作用、缓解水分亏缺、提高水分利用效率．而且促进植物的养分吸收、促进上层干土中分解、硝化和矿化等生化过程．尤其是发生在克隆植物植株间的水分共享这种特殊的水力提升,还影响群落的形成。然而,对于一般的水力提升而言,水力提升所释出水量到底有多少。其生态学作用有多大．有待深入研究。结合研究实践认为．要研究水力提升具有多大的生态学作用．必须准确地区分水;勺提升的水量和毛管水的上升水量。还建议了研究水力提升新的研究方法。