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。     

土壤-根系统水分再分配:土壤-植物-大气连续体中的一个小通路

吸收和传导水分一直被视为植物根系最主要的功能之一,而人们对根系在某些情况下还可以向土壤释放水分的事实及其对植物生长和生态系统功能的影响了解得还很不充分,尽管这样的证据由来已久。土壤-根系统水分再分配(Hydraulic redistribution, HR)是近20年间被发现和证实的,指水分从土壤中较湿的部分经由植物的根系传导而运动到土壤中较干的部分,通常发生在蒸腾减弱的夜间,可以沿水势梯度下降的方向而在不同土层间向上向下或侧向运动。HR研究揭示了土壤-植物-大气连续体中有时会存在土壤-根-土壤的水流小通路,细化了土壤-根系统中水分储存和运输的时空动态和机制。土壤水分状况的连续监测、根木质部液流测量、稳定性同位素技术的使用构成了HR实验研究的三大手段。当土壤中深层水分充足的时候,HR可以提高根系吸收和传导水分的效率,有利于植物充分利用资源,延长了浅层土壤的水分可利用期,有利于维持植物组织的生理活性和水流传导;旱季后降水来临的时候,HR可以将一部分降水转移到深层土壤,增加了可利用性水分的总量。对于干旱半干旱的沙地和草原、季节性干旱的森林等类型,HR过程可能对生态系统水分循环产生重要影响。有必要在国内针对这些生态系统展开深入的实验研究,同时探索将HR过程适当结合到生态系统模型和水文模型中,从而更准确地研究和预测群落内植物水分关系和生态系统水分动态。此外,结合农林设计、植被恢复、生态需水量估算和农业节水等方面进行的HR研究也值得深入探索。     

植物根-土界面水分再分配研究方法与影响因素

水分再分配是根-土界面根系对水分在垂直或水平方向的输导过程,对水分再分配的研究最早是在实验室盆栽条件下进行的,而盆栽测定的最基本方法就是分根法。实验室条件下,土壤水分的测定包括γ射线密度法、称量法、烘干法和时域反射仪（TDR）等。野外条件下,对根-土界面水分再分配的研究基本上有4种方法,利用热电隅干湿球温度计测定土壤水势的昼夜及季节波动、氢同位素示踪法、根木质部液流的测量和TDR测定土壤体积含水量。影响水分再分配的因素主要有土壤水分、土壤质地、植物种类及蒸腾作用等。在今后的研究中,应当加强对植物根-土界面水分再分配发生条件的研究,明确各影响因素与水分再分配发生的频率和数量的关系,从而为更好地利用其水分调节功能提供理论基础。     

植物根系水力再分配的研究进展

植物根系水力再分配(hydraulic redistribution,HR)是一种普遍现象,其提升数量大小与研究方法和影响因素有关.常用的研究方法包括土壤水分法、同位素示踪法、液流法、Ryel模型,每种方法均有缺点,单独使用其中任一方法其结果都存在差异.HR发生具有两个条件,即土壤水势(ψs)小于叶片水势(ψ1)且蒸腾...     

根-土界面水分再分配研究现状与展望

对根 -土界面水分再分配的研究背景 (概念、发现及证据 )、普遍性与再分配的水量及其生理生态学意义 (对相邻植物利用水分、根际活动、土壤 -植物 -大气系统水分传输和根系可塑性发育的促进效应 )等进行了深入论述 ,对水分再分配的认识和研究方法进行了探讨 ,并对未来的相关研究进行了展望     

植物水分利用策略研究进展

水分是影响植物生长发育的重要因子之一。地球上大多数生态系统中的植物都会经历一个降水相对稀少的干旱季节,植物在不同的季节与不同的生态系统中究竟如何利用水分,利用哪些水分去获得生存,成为一个人们关注的问题。在过去的20年,稳定同位素技术在植物生态学中的应用得到了稳定长足的发展。因为陆地植物(少数排盐种类除外)在水分吸收过程中不发生同位素分馏,因此可以利用δD与δ18O数据进行水分获取方式的研究。对植物木质部水分以及其潜在水源的稳定同位素进行分析,并参考土壤水势、叶片水势、土壤含水量等数据,同时运用二元或三元混合模型,可以定量确定植物的水分利用来源。大量的研究表明,不同功能型、生长阶段、季节的植物以及不同物种往往具有不同的水分利用策略。     

植物根系水力再分配模型参数分析与尺度转换

植物根系水力再分配(Hydraulic redistribution)是近几年提出的对植物根系水力提升现象一种更准确的描述。Ryel等(2002)建立的根系水力再分配模型(以下简称Ryel模型)模拟结果表明根系水力再分配是土壤水分动态的一个重要组成部分。该文基于Ryel模型,对模型中涉及的重要参数进行敏感性分析,更准确地阐述参数变化下根系水力再分配模型的行为动态,从而定量分析环境及植物自身等因素对根系水力再分配的影响。Ryel模型时间尺度和土层厚度的设定限制了模型的应用,该文通过参数调整,将模型从时间尺度为小时、土层厚度均一转换到时间尺度为天、土层厚度不等,并应用到内蒙古皇甫川流域。     

新疆艾比湖湿地自然保护区胡杨根系水分再分配的估算

根据对新疆艾比湖湿地自然保护区荒漠河岸林的主要建群种胡杨(Populus euphratica)的根系分布特征, 林冠下土壤的饱和容积含水量、最大导水率和根系最大导水率的试验观测数据, 以及对其林冠下5层不同深度土壤容积含水量、土壤水势的动态监测数据, 构建了胡杨根系水分再分配量估算的Ryel模型, 并对胡杨不同季节的水分再分配过程进行了短期模拟。结果表明: (1)胡杨根系水分再分配过程的水分再分配量的最大值出现在艾比湖地方时间凌晨2:30。(2)随着生长季节的变化, 胡杨根系水分再分配的作用逐渐减弱, 并表现出向土壤下层迁移的现象。6月份, 水分再分配过程主要发生在0-40 cm土层, 最大分配量为0.022 0 cm, 夜间总分配量为0.111 0 cm; 8月份水分再分配过程主要发生在10-70 cm土层, 最大分配量为0.006 5 cm, 夜间总分配量为0.018 4 cm; 10月份水分再分配过程主要发生在70-100 cm土层, 最大分配量为0.003 9 cm, 夜间总分配量为0.008 6 cm。     

土壤植物系统中植物根系吸收土壤水份研究进展

土壤植物系统是土壤—植物—大气连续体中的一个重要的子系统,该系统中的植物根系吸收土壤水份的研究已受到国内外的普遍重视,成为旱地农业生态系统中最为活跃的研究课题之一。从土壤—植物—大气连续体入手,对植物根系吸收土壤水份的影响因素,植物根系吸收土壤水份的微观模型及宏观模型等,进行了介绍和评述     

毛白杨根系液流与水力再分配特征

为确定毛白杨(Populus tomentosa)根系是否存在水力再分配现象,并探究其发生特征与影响因子,该研究以四年生毛白杨为研究对象,利用热比率法对3株样树的共计7条侧根(R1–R7)进行长期液流监测,并对土壤水分以及气象因子进行同步测定。结果显示:毛白杨存在两种水力再分配模式,分别为干旱驱动的水力提升和降雨驱动的水力下降,水力再分配的发生模式与特征受侧根分布深度与直径大小的影响。在整个生长季尺度上,毛白杨根系再分配的水量较低;但在极端干旱条件下,部分侧根再分配的水量可达其日总液流量的64.6%,表明水力再分配会为干旱侧根提供大量水分。根系吸水与气象-土壤的耦合因子(太阳辐射(R_s)×土壤含水率(SWC)、水汽压亏缺(VPD)×SWC、参考蒸散发(ET_o)×SWC)间存在显著相关关系,但水力再分配与所选因子基本不相关。此外,毛白杨浅层根中存在特殊的日间逆向液流现象,其液流量最高可占日液流总量的79.2%(R1)到90.7%(R2),该现象可能对浅层根系抗旱起到重要作用。     

Hydraulic redistribution in three Amazonian trees

About half of the Amazon rainforest is subject to seasonal droughts of 3 months or more. Despite this drought, several studies have shown that these forests, under a strongly seasonal climate, do not exhibit significant water stress during the dry season. In addition to deep soil water uptake, another contributing explanation for the absence of plant water stress during drought is the process of hydraulic redistribution; the nocturnal transfer of water by roots from moist to dry regions of the soil profile. Here, we present data on patterns of soil moisture and sap flow in roots of three dimorphic-rooted species in the Tapajós Forest, Amazônia, which demonstrate both upward (hydraulic lift) and downward hydraulic redistribution. We measured sap flow in lateral and tap roots of our three study species over a 2-year period using the heat ratio method, a sap-flow technique that allows bi-directional measurement of water flow. On certain nights during the dry season, reverse or acropetal flow (i.e.,in the direction of the soil) in the lateral roots and positive or basipetal sap flow (toward the plant) in the tap roots of Coussarea racemosa (caferana), Manilkara huberi (maçaranduba) and Protium robustum (breu) were observed, a pattern consistent with upward hydraulic redistribution (hydraulic lift). With the onset of heavy rains, this pattern reversed, with continuous night-time acropetal sap flow in the tap root and basipetal sap flow in lateral roots, indicating water movement from wet top soil to dry deeper soils (downward hydraulic redistribution). Both patterns were present in trees within a rainfall exclusion plot (Seca Floresta) and to a more limited extent in the control plot. Although hydraulic redistribution has traditionally been associated with arid or strongly seasonal environments, our findings now suggest that it is important in ameliorating water stress and improving rain infiltration in Amazonian rainforests. This has broad implications for understanding and modeling ecosystem process and forest function in this important biome.     

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.     

Hydraulic redistribution of soil water in two old-growth coniferous forests: quantifying patterns and controls

Although hydraulic redistribution of soil water (HR) by roots is a widespread phenomenon, the processes governing spatial and temporal patterns of HR are not well understood. We incorporated soil/plant biophysical properties into a simple model based on Darcy's law to predict seasonal trajectories of HR. We investigated the spatial and temporal variability of HR across multiple years in two old-growth coniferous forest ecosystems with contrasting species and moisture regimes by measurement of soil water content (theta) and water potential (Psi) throughout the upper soil profile, root distribution and conductivity, and relevant climate variables. Large HR variability within sites (0-0.5 mm d(-1)) was attributed to spatial patterns of roots, soil moisture and depletion. HR accounted for 3-9% of estimated total site water depletion seasonally, peaking at 0.16 mm d(-1) (ponderosa pine; Pinus ponderosa) or 0.30 mm d(-1) (Douglas-fir; Pseudotsuga menziesii), then declining as modeled pathway conductance dropped with increasing root cavitation. While HR can vary tremendously within a site, among years and among ecosystems, this variability can be explained by natural variability in Psi gradients and seasonal courses of root conductivity.     

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.     

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.     

Hydraulic redistribution in a Douglas-fir forest: lessons from system manipulations

Hydraulic redistribution (HR) occurs in many ecosystems; however, key questions remain about its consequences at the ecosystem level. The objectives of the present study were to quantify seasonal variation in HR and its driving force, and to manipulate the soil-root system to elucidate physiological components controlling HR and utilization of redistributed water. In the upper soil layer of a young Douglas-fir forest, HR was negligible in early summer, but increased to 0.17 mm day(-1) (20-60 cm layer) by late August when soil water potential was approximately -1 MPa. When maximum HR rates were observed, redistributed water replenished approximately 40% of the water depleted from the upper soil on a daily basis. Manipulations to the soil or to the soil/plant water potential driving force altered the rate of observed HR indicating that the rate of HR is controlled by a complex interplay between competing soil and plant water potential gradients and pathway resistances. Separating roots from the transpiring tree resulted in increased HR, and sap flow measurements on connected and disconnected roots showed reversal of water flow, a prerequisite for HR. Irrigating a small plot with deuterated water demonstrated that redistributed water was taken up by small understorey plants as far as 5 m from the watering source, and potentially further, but the utilization pattern was patchy. HR in the upper soil layers near the watering plot was twice that of the control HR. This increase in HR also increased the amount of water utilized by plants from the upper soil. These results indicate that the seasonal timing and magnitude of HR was strongly governed by the development of water potential differences within the soil, and the competing demand for water by the above ground portion of the tree.     

Temporal variation in plant-soil feedback controls succession

Soil abiotic and biotic factors play key roles in plant community dynamics. However, little is known about how soil biota influence vegetation changes over time. Here, we show that the effects of soil organisms may depend on both the successional development of ecosystems and on the successional position of the plants involved. In model systems of plants and soils from different successional stages, we observed negative plant–soil feedback for early-successional plant species, neutral feedback for mid-successional species, and positive feedback for late-successional species. The negative feedback of early-successional plants was independent of soil origin, while late-successional plants performed best in late- and worst in early-successional soil. Increased performance of the subordinate, late-successional plants resulted in enhanced plant community diversity. Observed feedback effects were more related to soil biota than to abiotic conditions. Our results show that temporal variations in plant–soil interactions profoundly contribute to plant community assemblage and ecosystem development.