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.     

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.     

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。     

Depressed hydraulic redistribution of roots more by stem refilling than by nocturnal transpiration for Populus euphratica Oliv. in situ measurement

During the night, plant water loss can occur either through the roots, as hydraulic redistribution (HR), or through the leaves via the stoma, as nocturnal transpiration (E_n), which was methodologically difficult to separate from stem refilling (R_e). While HR and E_n have been reported across a range of species, ecosystem, and climate zone, there is little understanding on the interactions between E_n and/or R_e and HR. As water movement at night occurs via gradients of water potential, it is expected that during periods of high atmospheric vapor pressure deficit (VPD), water loss via E_n will override water loss via HR. To test this hypothesis, sap flow in stems and roots of Populus euphratica Oliv. trees, growing in a riparian zone in a hyperarid climate, was measured once in a year. Nocturnal stem sap flow was separated into E_n and R_e using the “forecasted refilling” method. Substantial nocturnal sap flow (38% of 24‐hr flux on average) was observed and positively correlated with VPD; however, the strength of the correlation was lower (R² = .55) than diurnal sap flow (E_d) (R² = .72), suggesting that nocturnal stem sap flow was attributed to both water loss through the canopy and replenishment of water in stem tissues. Partitioning of nocturnal sap flow shows that R_e constituted approximately 80%, and E_n ~20%, of nocturnal sap flow. The amount of root sap flow attributed to redistribution was negatively related to E_d (R² = .69) and the amount of acropetally sap flow in stems, R_e (R² = .41) and E_n (R² = .14). It was suggested that the magnitude of HR is more strongly depressed by R_e that was recharge to the water loss via E_d than by E_n. It was consistent with whole‐tree water balance theory, that the nighttime upward sap flow to xylem, stem refilling and transpiration, may depress hydraulic redistribution of roots.     

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。     

Phylogenetic and ecological patterns in nighttime transpiration among five members of the genus Rubus co‐occurring in western Oregon

Nighttime transpiration is a substantial portion of ecosystem water budgets, but few studies compare water use of closely related co‐occurring species in a phylogenetic context. Nighttime transpiration can range up to 69% of daytime rates and vary between species, ecosystem, and functional type. We examined leaf‐level daytime and nighttime gas exchange of five species of the genus Rubus co‐occurring in the Pacific Northwest of western North America in a greenhouse common garden. Contrary to expectations, nighttime transpiration was not correlated to daytime water use. Nighttime transpiration showed pronounced phylogenetic signals, but the proportion of variation explained by different phylogenetic groupings varied across datasets. Leaf osmotic water potential, water potential at turgor loss point, stomatal size, and specific leaf area were correlated with phylogeny but did not readily explain variation in nighttime transpiration. Patterns in interspecific variation as well as a disconnect between rates of daytime and nighttime transpiration suggest that variation in nighttime water use may be at least partly driven by genetic factors independent of those that control daytime water use. Future work with co‐occurring congeneric systems is needed to establish the generality of these results and may help determine the mechanism driving interspecific variation in nighttime water use.     

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     

植物根水倒流的证据及意义

对根水倒流的研究进展进行了综述,分析和介绍了其存在的类型、普遍性、发生量、发生位置、生理学基础等,讨论了其在植物生理学、生态学、营养学和作物载培学等学科中的重要意义,并是其存在的问题和进一步研究的方向。     

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.     

Seasonal changes of root hydraulic conductance (KRL) in four forest trees: an ecological interpretation

This paper deals with the possibility of relating root hydraulic parameters to an ecological index describing the continentality/oceanicity of four forest trees. Root hydraulic conductance ($K_R$) of seedlings of Fagus sylvatica L., Quercus ilex L., Quercus suber L. and Quercus pubescens Willd. was measured in May, August and November 1996. $K_R$ was calculated in terms of the relation of the water flow through intact root systems in situ measured with the pressure chamber, and the pressure driving the flow. The sufficiency of the root system to supply the foliage was estimated by dividing $K_R$ by the seedlings leaf surface area ($A_L$) thus obtaining $K_RL$. In the spring, $K_RL$ was largest in F. sylvatica and smallest in Q. pubescens with intermediate values recorded in Q. ilex and Q. suber. All the species studied showed a large decline in $K_RL$ just prior to the winter rest except for Q. suber which mantained $K_RL$ approximately constant through the period of study. In most cases, $K_RL$ changed in accordance with analogous changes in the flow. When the total seedlings' leaf surface area ($A_L$) was plotted versus $K_RL$, it appeared that $K_RL$ of Q. pubescens increased with $A_L$, proportionally, while $K_RL$ of F. sylvatica was inversely related to $A_L$. This, together with the largest $K_RL$ recorded in the summer in Q. pubescens, was interpreted as advantageous to this species (which is adapted to semi-arid environments) in that: (a) roots could supply water to foliage efficiently even during the adverse season and (b) the foliage growth could be sustained even in summer.No statistically significant relation of $K_RL$ to the continentality index calculated for the four species studied on the basis of their European distribution, was found to exist. Nonetheless, our data appear to be encouraging for future research aimed at better interpreting the typical distribution areas of plant species.     

Drought resistance of Quercus pubescens as a function of root hydraulic conductance, xylem embolism and hydraulic architecture

Water relations, xylem embolism, root and shoot hydraulic conductance of both young plants in the field and potted seedlings of Quercus pubescens have been studied with the aim of investigating whether these variables may account for the well known adaptation of this oak species to arid habitats. Our data revealed that Q. pubescens is able to maintain high leaf relative water contents under water stress conditions. In fact, relative water contents measured in summer (July) did not differ from those recorded in April. This was apparently achieved by compensating water loss by an equal amount of water uptake. Such a drought avoidance strategy was made possible by the recorded high hydraulic efficiency of stems and roots under water stress. In fact, root hydraulic conductance of field-grown plants was maintained high in summer when the percentage loss of hydraulic conductance of stems was lowest. The hydraulic architecture of young plants of Q. pubescens measured in terms of partitioning of hydraulic resistances along the water pathway revealed that the highest hydraulic resistance was located in stems of the current year's growth. This hydraulic architecture is interpreted as consistent with the adaptation of Q. pubescens to arid habitats as a consequence of the recorded seasonal changes in water relation parameters as well as in root and stem hydraulics.     

Seasonal variation in native hydraulic conductivity between two deciduous oak species

植物抗旱性机制包括耐旱性和避旱性。耐旱性的特征主要是木质部易栓塞性和膨压损失点,而避旱性的特征则是叶片脱落和落叶性。虽然这些机制可以权衡物种连续性,但对于干旱引发的叶片脱落和持续时间的变化如何影响木质部和叶片耐受性之间的关系却知之甚少。在本研究中,我们研究两种具有不同叶片脱落叶行为的橡树(栎属)的耐旱性程度差异。我们预测了Quercus deserticola 在旱季结束时落叶(晚落叶)而面临更大的空化风险,导致其比Q. laeta更耐旱,对水分的利用也更加保守。Q. laeta 只在旱季中期很短的时间内落叶(短落叶)。本研究在墨西哥中部进行,两种橡树各具有单个种群,它们之间相距1.58公里,海拔差为191米。Q. deserticola (晚落叶)多出现在下坡,而Q. laeta (短落叶)多出现在沿坡度海拔较高的地方。我们评估了样地内茎干的水分传导率的季节差异(雨季和旱季),并检测了两个物种间木质部对空化的脆弱性、叶片水分利用和叶片膨松损失点的差异。两种橡树在耐旱性状方面没有显著差异,主要包括木质部易栓塞性、叶片膨松损失点和气孔导度。然而,两种植物在旱季都有不同的表现：短落叶植物对木质部功能的负面影响要比晚落叶植物低。综上所述,两种橡树在植物生理性能方面的季节变化取决于冠层叶面积的减少。     

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。     

整树水力导度协同冠层气孔导度调节森林蒸腾

冠层气孔导度决定森林的蒸腾效率,它对驱动水汽移动的水汽应力的响应受树木水力结构的影响,并随水汽压亏缺上升和水力导度下降而降低,维持水势在最低阈值之上,避免出现水力灾变,调控冠层蒸腾。由于叶形和树冠结构的特点,部分脱耦联反映了湿润地区阔叶林冠层与大气的水汽交换特征,单纯以气孔导度的变化难以完整描述水分通量的调节规律,因而,需要考虑冠层气孔导度与水力导度协同控制冠层蒸腾的潜在机理。通过整合叶片气孔气体交换、树干液流、冠层微气象和其他环境因子的野外观测值,估测不同时间尺度的森林冠层气孔导度与大气的脱耦联系数和变异范围,以基于树干液流的冠层蒸腾,结合叶片/土壤水势梯度计算的水力导度,分析水力导度影响冠层气孔导度响应水汽压亏缺的敏感性,可以揭示和阐明水力导度和冠层气孔导度联合调节森林蒸腾的机理,对准确估测全球变化背景下森林对水资源利用的潜在生态效应有明显的理论意义。     

Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.     

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.     

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2–4-mm diameter) collected at 20–30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance (g_s) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20–30 cm depth above –0.5 MPa in the old-growth Douglas-fir site and –1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about –2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g_s decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.     

The spittlebug Philaenus spumarius feeds from mature xylem at the full hydraulic tension of the transpiration stream

Feeding dynamics in the spittlebug Philaenus spumarius were analysed to determine the food source. Evidence from previous reports indicates that this species and similar insects feed from the xylem. However, it is not clear whether they extract sap from the main transpiration stream, or from some subsidiary compartment of xylem, such as immature or cavitated vessels. In feeding from such subsidiary compartments, the insects could avoid the large hydraulic tensions thought to prevail in xylem on the main transpiration stream. Philaenus , for example, is shown here to initiate feeding, and to feed rapidly, from plants in which the full xylem tension is estimated at −10 bar. From measurements of feeding rates and feeding patterns in stressed and unstressed plants, it is shown that Philaenus cannot feed from subsidiary xylem compartments. It is concluded that this insect feeds directly from the main transpiration stream, and at the full hydraulic tension which prevails there. The anatomy of the feeding pump in Philaenus appears consistent with this conclusion. Spittlebugs therefore offer an experimental tool for quasi- non-invasive sampling of the transpiration stream.     

K+ status and ABA affect both exudation rate and hydraulic conductivity in sunflower roots

Twenty‐day‐old sunflower plants ( Helianthus annuus L. cv. Sun‐Gro 380) grown in nutrient solutions with different KCl levels were used to study the effects of K⁺ status of the root and of abcisic acid (ABA) on the exudation rate (J_v), the hydraulic conductivity of the root (L_p), the fluxes of exuded K⁺ and Na⁺ (J_K and J_Na), and the gradient of osmotic pressure between the xylem and the external medium. J_v and L_p increased in direct proportion to the K⁺ starvation of the root. Also addition of ABA (4 µ M ) at the onset of exudation in the external medium made J_v and L_p rise, and this effect also increased with the degree of K⁺ starvation. Similarly, K⁺ starvation and ABA promoted both the flux of exuded Na⁺ and the accumulation of Na⁺ in the root. We suggest that ABA acts as a regulating signal for the radial transport of water across the root, and that potassium may be an effector of this mechanism.