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.     

The relationship of abscisic acid metabolism to stomatal conductance in Douglas-fir during water stress

Changes in abscisic acid and its metabolites were followed through two drought cycles in Pseudotsuga menziesii (Mirb.) Franco seedlings to determine the metabolic pathway of the hormone and its relationship to branch (stomatal) conductance. Three year-old, intact seedlings were water-stressed, watered, and restressed over a period of 30 days. Water potential was sampled with a pressure chamber and branch conductance with a steady-state porometer. Needle content of abscisic acid and 2- trans -abscisic acid and their saponifiable conjugates were quantified with gas-liquid chromatography. The typical water potential threshold in branch conductance, decreasing abruptly at -2.0 MPa, corresponded to an increase in abscisic acid content of 240 ng g⁻¹. The relationship between abscisic acid and water potential was not definitive, though the general trend was an increase in the hormone with intensifying stress until water potential was -5.0 MPa, when concentration sharply declined. No adjustment to stress was observed in the relationships, but stress during the second cycle progressed more slowly. A linear relationship between abscisic acid and its conjugate indicated the importance of the interconversion of the two compounds for storage and supply of the free acid.     

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.     

Impacts of tree height on leaf hydraulic architecture and stomatal control in Douglas-fir

This study investigated the mechanisms involved in the regulation of stomatal closure in Douglas-fir and evaluated the potential impact of compensatory adjustments in response to increasing tree height upon these mechanisms. In the laboratory, we measured leaf hydraulic conductance (K(leaf)) as leaf water potential (Psi(l)) declined for comparison with in situ diurnal patterns of stomatal conductance (g(s)) and Psi(l) in Douglas-fir across a height gradient, allowing us to infer linkages between diurnal changes in K(leaf) and g(s). A recently developed timed rehydration technique was used in conjunction with data from pressure-volume curves to develop hydraulic vulnerability curves for needles attached to small twigs. Laboratory-measured K(leaf) declined with increasing leaf water stress and was substantially reduced at Psi(l) values of -1.34, -1.45, -1.56 and -1.92 MPa for foliage sampled at mean heights of approximately 20, 35, 44 and 55 m, respectively. In situ g(s) measurements showed that stomatal closure was initiated at Psi(l) values of -1.21, -1.36, -1.74 and -1.86 MPa along the height gradient, which was highly correlated with Psi(l) values at loss of K(leaf). Cryogenic scanning electron microscopy (SEM) images showed that relative abundances of embolized tracheids in the central vein increased with increasing leaf water stress. Leaf embolism appeared to be coupled to changes in g(s) and might perform a vital function in stomatal regulation of plant water status and water transport in conifers. The observed trends in g(s) and K(leaf) in response to changes in Psi(l) along a height gradient suggest that the foliage at the tops of tall trees is capable of maintaining stomatal conductance at more negative Psi(l). This adaptation may allow taller trees to continue to photosynthesize during periods of greater water stress.     

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 of water from Pinus ponderosa trees to seedlings: evidence for an ectomycorrhizal pathway

While there is strong evidence for hydraulic redistribution (HR) of soil water by trees, it is not known if common mycorrhizal networks (CMN) can facilitate HR from mature trees to seedlings under field conditions. Ponderosa pine (Pinus ponderosa) seedlings were planted into root-excluding 61-microm mesh barrier chambers buried in an old-growth pine forest. After 2 yr, several mature trees were cut and water enriched in D(2)O and acid fuchsin dye was applied to the stumps. Fine roots and mycorrhizal root tips of source trees became heavily dyed, indicating reverse sap flow in root xylem transported water from stems throughout root systems to the root hyphal mantle that interfaces with CMN. Within 3 d, D(2)O was found in mesh-chamber seedling foliage > 1 m from source trees; after 3 wk, eight of 10 mesh-chamber seedling stem samples were significantly enriched above background levels. Average mesh-chamber enrichment was 1.8 x greater than that for two seedlings for which the connections to CMN were broken by trenching before D(2)O application. Even small amounts of water provided to mycorrhizas by HR may maintain hyphal viability and facilitate nutrient uptake under drying conditions, which may provide an advantage to seedlings hydraulically linked by CMN to large trees.     

Water stress, shoot growth and storage of non-structural carbohydrates along a tree height gradient in a tall conifer

We analysed concentrations of starch, sucrose, glucose and fructose in upper branch wood, foliage and trunk sapwood of Douglas-fir trees in height classes ranging from ~2 to ~57 m. Mean concentrations of non-structural carbohydrates (NSC) for all tissues were highest in the tallest height class and lowest in the lowest height class, and height-related trends in NSC were most pronounced in branches. Throughout a 17-month sampling period, mean values of branch NSC from the 57 m trees ranged between 30 and 377% greater than the 2 m trees. Branch NSC was inversely correlated with midday shoot water potential (Ψ(l)), shoot osmotic potential at full turgor (Ψ) and shoot extension. Temporal fluctuation in branch NSC was inversely correlated with height, and positively correlated with midday Ψ(l) , Ψ and shoot extension. The positive correlation between height and storage of NSC, and the negative correlation between NSC storage and shoot extension provide evidence that size-related growth decline in trees is not strongly associated with constraints on photosynthesis. The negative correlation between height and fluctuation in NSC suggests that mobilization of photosynthate in taller trees is constrained by some factor such as reductions in turgor-driven cell expansion or constraints on phloem transport.     

Douglas-fir root biomass and rooting profile in relation to soils in a mid-elevation area (Beaujolais Mounts,France)

Douglas-fir is the main reforestation species in the French Massif Central area (14 000 ha), but little is known about its rooting strategy in different soil conditions. This information has important implications for the choice of better soils for settling Douglas-fir, and consequently limiting risks of failure, pests or diseases. As a result, the influence of edaphic conditions on rooting patterns of dominant Douglas-fir was studied over a large range of ecological conditions in a mid-elevation area of the French Massif Central (Beaujolais Mounts). Root systems were studied extensively using the trench profile wall technique and the sector method in 74 pure and evenly aged Douglas-fir stands. The stands were chosen as being representative of soil conditions among 165 stands in an auto-ecological study. The rooting patterns were related to seven typical soil profiles, and to root profile groups. Results stressed that edaphic constraints due to substratum and soil structures have a strong influence on root system morphology. Important variations in root biomass and vertical distribution were highlighted among soils. Small fine root biomass is maximal for soils with no major edaphic constraints. The vertical distribution of fine root biomass is positively correlated for some soil types with organic C, total N, and most cations. For some types it was negatively correlated with the amount of exchangeable aluminum and coarse fragments, and with constraining rock facies. Harsher soils however, showed no correlation between soil chemical variables and fine-root biomass. A practical implication is that Douglas-fir seems to be a pliable and adaptive species: variation in habit and root system biomass are considerable within a study area which was presumed uniform.     

Hydraulic control of stomatal conductance in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and alder [Alnus rubra (Bong)] seedlings

Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (g_c). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in g_c with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.     

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。     

Hydraulic lift in drought-tolerant and -susceptible maize hybrids

Hydraulic lift was investigated in a greenhouse study involving two drought-tolerant maize (Zea mays L.) hybrids (TAES176 and P3223) and a drought-susceptible hybrid (P3225) during the flowering stage. Root systems were grown in two soil compartments – a drier upper soil and a wetter deep soil. The plants were shaded for 3 h during the daytime. Soil volumetric water content (Ø_v) in the upper pots was measured with time domain reflectometry (TDR) before and after shading. An increase in Ø_v in the upper pot was detected with TDR in the drought-tolerant hybrids following 3 h of shading, but not in the drought-susceptible hybrid. Furthermore, water exuded from roots in the top soil layers was greater in the more drought-tolerant TAES176 than in P3223 (489 vs. 288 g per pot in 3 h, P<0.005). The sizable amount of water from hydraulic lift allowed TAES176 to reach a peak transpiration rate 27–42% higher than the drought-susceptible hybrid P3225 on the days when the evaporative demand was high. To our knowledge, this is the first experiment that reveals a significant surge of transpiration due to hydraulic lift following midday shading. Hydraulic lift also prevented soil moisture depletion in the upper pots with TAES176, but not with P3223 or P3225. Root characteristics may be responsible for differences in hydraulic lift of the three maize hybrids. There were 2.3–3.3-fold more primary roots in the deep moist soil in P3223 and TAES176 than in P3225 that may enable these hybrids to absorb and transport water at faster rates. Therefore, more water can be exuded into the upper drier soil when transpiration is suppressed by shading. Larger primary roots (20–28% larger diameter) and a higher root volume in the upper soil in TAES176 and P3223 than in P3225 may contribute to higher root hydraulic conductance and greater water efflux from the roots. The negligible hydraulic lift in P3225 may also relate to higher night-time transpiration of the hybrid. This report has documented, for the first time, the existence of genetic variations in hydraulic lift among maize hybrids and links between hydraulic lift and drought tolerance within maize plants. It appears that one of drought tolerance mechanisms in maize may lie in the extent of hydraulic lift.     

Leaf water 18O and 2H enrichment along vertical canopy profiles in a broadleaved and a conifer forest tree

Distinguishing meteorological and plant‐mediated drivers of leaf water isotopic enrichment is prerequisite for ecological interpretations of stable hydrogen and oxygen isotopes in plant tissue. We measured input and leaf water δ²H and δ¹⁸O as well as micrometeorological and leaf morpho‐physiological variables along a vertical gradient in a mature angiosperm (European beech) and gymnosperm (Douglas fir) tree. We used these variables and different enrichment models to quantify the influence of Péclet and non‐steady state effects and of the biophysical drivers on leaf water enrichment. The two‐pool model accurately described the diurnal variation of leaf water enrichment. The estimated unenriched water fraction was linked to leaf dry matter content across the canopy heights. Non‐steady state effects and reduced stomatal conductance caused a higher enrichment of Douglas fir compared to beech leaf water. A dynamic effect analyses revealed that the light‐induced vertical gradients of stomatal conductance and leaf temperature outbalanced each other in their effects on evaporative enrichment. We conclude that neither vertical canopy gradients nor the Péclet effect is important for estimates and interpretation of isotopic leaf water enrichment in hypostomatous trees. Contrarily, species‐specific non‐steady state effects and leaf temperatures as well as the water vapour isotope composition need careful consideration.     

亚热带森林演替树种叶片气孔导度对环境水分的水力响应

利用LI-1600稳态气孔计和PMS压力室,在田间测定了群落演替早期强阳生性树种桃金娘(Rhodomyrtus tomentosa)和三叉苦(Evodia lepta)、偏中性的阳生性树种荷木(Schima superba)、群落演替后期的耐荫树种鸭脚木(Schefllera octophylla)和九节(Psychotrie rubra)的叶片气孔导度(gs)和叶片水势(ΨL),研究不同演替阶段树种的气孔导度对环境水分的响应.结果表明,早上叶片有较高的ΨL,随着时间推移ΨL逐渐降低,与此同时比叶水力导度(KL)随ΨL降低而下降,桃金娘、三叉苦、荷木、鸭脚木和九节水力导度初始最低值时的ΨL分别为-1.6、-1.42、-1.30、-0.9MPa和-1.05MPa.随着ΨL降低,田间测定的gs开始从上午的较低值上升至约中午时的最大值,随后开始降低,此时的ΨL分别为-1.58、-1.52、-1.35、-1.02MPa和-1.0MPa.不同植物种类有不同的KL初始最低值的ΨL和gs达到最大值的ΨL.但不论何种供试树种,KL最低值时的ΨL与gs开始从最大值下降时的ΨL相近;显示KL与gs在动态变化中存在协调关系.树种间的gs和KL对ΨL的不同响应显示桃金娘和三叉苦的KL最低值时和gs开始下降时的ΨL均较鸭脚木和九节对应的ΨL低(p<0.05),意味着演替早期树种能在较强水分胁迫下保持较高的气孔导度.这一水力特性保证树种在水分胁迫下维持叶片的光合速率,有利于其在群落中的生长和优势地位的维护,而演替后期树种在较高ΨL下气孔关闭,降低了光合速率.全球变暖和环境进一步干旱可能成为限制亚热带森林植物群落的正向演替进程的潜在因素之一.     

坡地赤红壤—芒果—大气连续体水分运移力能关系的田间试验研究

依据田间实测资料,运用水流的电模拟原理,定量分析了坡地赤红壤—芒果—大气连续体水分运移过程中水分势能和水流阻力各分量的大小、变化规律及其相对重要性．结果表明,水流在该连续体中运移时,其能量主要消耗在由叶部到大气这一环节上,水分势能损失达１５．４４９１～５８．４０３６ＭＰａ,水分流经土壤到达叶部,水分热能降低０．２８１８～０．６９７９ＭＰａ;水流阻力主要分布于从叶气孔腔到大气的扩散过程,叶—气系统的水流阻力占该连续体水流总阻力的９５．６８％～９９．０３％,植株体的水流阻力占该连续体水流总阻力的０．９７％～４．３２％,而土壤阻力相对可忽略不计．     

Use of a microtensiometer technique to study hydraulic lift in a sandy soil planted with pearl millet (Pennisetum americanum [L.] Leeke)

A pot experiment was carried out with pearl millet (Pennisetum americanum [L.] Leeke) growing in a sandy soil in which the upper (topsoil) and lower (subsoil) parts of the pots were separated by a perlite layer to prevent capillary water movement. Using microtensiometers a study was made to establish whether it was possible to measure hydraulic lift by which the upper part of the soil was rewetted when water was supplied exclusively to the lower part of the soil.Hydraulic lift occurred during the first seven days of the period of measurement, with a maximum water release to the soil of 2.7 Vol. % during one night (equivalent to 10.8 mL water in the top 10 cm of the soil profile). This magnitude was obtained at very high root length densities, so that water release from the roots would be expected to be much smaller under field conditions.Hydraulic lift ceased when the soil matric potential in the topsoil dropped below-10 kPa at the end of the light period and could not be re-established, neither by extending the dark period, nor after rewatering the topsoil. The disappearance of hydraulic lift could be explained in part through osmotic adaptation of plant roots and, thus prevention of water release from the roots in the topsoil. It is concluded that hydraulic lift may affect nutrient uptake from drying topsoil by extending the time period favourable for uptake from the topsoil.     

次生栎林与火炬松人工林土壤呼吸的季节变异及其主要影响因子

土壤呼吸是陆地生态系统碳循环的重要组成部分.随着全球气候变暖趋势逐渐明显,土壤呼吸的时空变异及其对温度变化的响应已成为生态学研究的重要内容之一.利用LI-6400-09土壤碳通量观测仪,在江苏省南京林业大学下蜀实验基地,采用随机区组实验设计方法,连续两年测定了北亚热带次生栎林和火炬松人工林土壤呼吸的季节动态变化,结果表明:(1)两种林分内土壤呼吸速率均具有明显的季节波动,表现为:在最冷的1月份,土壤呼吸速率最低,随着土壤温度的升高,土壤呼吸速率也逐渐上升,在7、8月份达到最大值,随后又逐渐下降;(2)次生栎林月平均土壤呼吸速率在0.271～3.22μmolCO2 · m-2 · s-1之间,年变异幅度为11.88;火炬松人工林月平均土壤呼吸速率在0.336～3.06μmolCO2 · m-2 · s-1 ,年变异幅度为9.11;(3)次生栎林土壤呼吸的 Q10值在2.19至2.27之间,火炬松人工林土壤呼吸的Q10值在2.02至2.15之间,次生栎林土壤呼吸对温度的敏感性大于火炬松人工林;(4)土壤呼吸速率与不同深度层次土壤温度之间均呈显著性正相关,与土壤微生物生物量之间呈显著性负相关,而与土壤含水率、凋落物输入量之间相关不显著.研究结果初步阐明了江淮流域北亚热带典型森林植被土壤呼吸的季节动态特征及主要影响因子,为进一步揭示该区域森林土壤碳循环特点提供了理论基础.     

Hydraulic Lift in Cork Oak Trees in a Savannah-Type Mediterranean Ecosystem and its Contribution to the Local Water Balance

The aim of this study was to identify the sources and depth of water uptake by 15-years old Quercus suber L. trees in southern Portugal under a Mediterranean climate, measuring δ¹⁸O and δD in the soil–plant-atmosphere continuum. Evidence for hydraulic lift was substantiated by the daily fluctuations observed in Ψ_s at 0.4 and 1 m depth and supported by similar δ¹⁸O values found in tree xylem sap, soil water in the rhizosphere and groundwater. From 0.25 m down to a depth of 1 m, δD trends differed according to vegetation type, showing a more depleted value in soil water collected under the evergreen trees (−47‰) than under dead grasses (−35‰). The hypothesis of a fractionation process occurring in the soil due to diffusion of water vapour in the dry soil is proposed to explain the more depleted soil δD signature observed under trees. Hydraulically lifted water was estimated to account for 17–81% of the water used during the following day by tree transpiration at the peak of the drought season, i.e., 0.1–14 L tree⁻¹ day⁻¹. Significant relationships found between xylem sap isotopic composition and leaf water potential in early September emphasized the positive impact of the redistribution of groundwater in the rhizosphere on tree water status.