Water transport through plant tissue: the apoplasm and symplasm pathways

A detailed quantitative analysis of water flow through the apoplasm and symplasm of plant tissue is presented. The analysis results in two coupled diffusion equations which describe water transport in the two pathways. Various parameters entering the analysis identify the physical properties of the tissue which control the transport process as the resistance to water flow per cell in the two parallel pathways, the resistance per cell between pathways, and the water capacity per cell in the two pathways. Values for the several resistances and water capacities are estimated from available data, and a model problem is solved wherein a sheet of tissue at an initial water potential of — δ bars is immersed in a container of water. The resulting solutions show that depending on the values assigned to the controlling parameters, local water potential equilibrium between each cell and its cell wall may or may not obtain. In the special case of local equilibrium (water potential in the symplasm and apoplasm pathways essentially equal), the transport process can be described by a single diffusion equation which is derived along with an expression for the tissue diffusivity. It is concluded that the weakest link in the analysis is the estimated value for the permeability of the plasmodesma membrane, and that a logical extension of the theory would be to include the effects of a diffusable solute.     

Water Flow in Beta vulgaris Storage Tissue

The relative magnitudes of the hydraulic resistances, water capacities, and water potential equilibration time constants for the single cell, for the apoplast, and for the symplast in higher plant tissue are assessed. Swelling of beetroot (Beta vulgaris, var. `Detroit Red') storage tissue sections in pure water is measured using a displacement transducer. This method of measurement avoids the difficulty of solute diffusion in the apoplast. Theoretical analysis of the experimental results shows that the main path of water flow into the tissue is the apoplast rather than the symplast, that the main resistance to water flow into the cells is usually the cell membrane rather than the apoplast, but that in some cases the apoplast resistance and water capacity can contribute significantly to the water potential equilibration time constant of the tissue.     

Transport of water and solutes across bullfrog alveolar epithelium

Water and solute transport properties of the alveolar epithelium of isolated bullfrog lungs were studied. Lungs from Rana catesbeiana were removed and mounted in an Ussing chamber. Unstirred layers on both sides of the tissue were estimated from the time courses of dilution potential development, and the measured transport parameters were corrected for the effect of the unstirred layers. Spontaneous potential difference, short-circuit current, tissue resistance, instantaneous voltage-current relationships, diffusional permeabilities of water and hydrophilic solutes, and hydraulic conductivities were determined. The hydraulic conductivity obtained from hydrostatically driven water flow anomalously decreased with time, and was initially 100 -1,000 times higher than osmotically determined hydraulic conductivity. The equivalent pore radius of the bullfrog alveolar epithelium was estimated to be 0.8-0.9 nm. We conclude that the alveolar epithelium is extremely tight, presenting a major barrier to water and solute flow. This high resistance to water and solute flow may be helpful in maintaining the alveolar lumen relatively free of fluid under normal physiological conditions.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF COSTA RICAN BLECHNUM FERNS

Hydraulic conductivities of stems, stipes, and elongate leaf stipes were determined for greenhouse-grown Blechnum (B. fraxineum, B. fragile, B. buchtienii, B. sprucei) and Salpichlaena (S. volubilis) plants collected in tropical rain forests of Costa Rica. Organ conductivity was examined in relation to morphology and tracheid characteristics in order to gain an understanding of factors influencing water flow. Hydraulic conductivity of plant organs was determined by measurement of transpiration rates, leaf areas, and water potential gradients. Erect stemmed species develop larger whole plant water potential gradients than elongate stemmed species for a similar transpiration rate. Elongate leaves develop even smaller water potential gradients for a given transpiration rate. Stems have larger hydraulic conductivities but smaller leaf-specific conductivities (LSCs) than stipes. Small conductivities and small LSCs are associated with short, erect stems. Elongate structures have large conductivities and large LSCs. Of the tracheid characteristics examined, the most important characteristics determining the magnitude of organ hydraulic conductivity are diameter, pit aperture area between tracheids, taper length, and cell length. Large conductivities of S. volubilis climbing leaf stipes are associated with very large-diameter tracheids (some > 200 μm), large tracheid number, exceptionally long tracheids (some > 4 cm), large pit aperture area between tracheids, short tracheid taper, and smooth tracheid lumen walls. Hagen-Poiseuille estimates of hydraulic conductivity range from 1.1 to 3.3 times the measured values. Conductivity of stipes is highly correlated with leaf area supplied by stipes. Conductivities of stems and elongate leaf stipes also correlate with leaf area supplied by these structures. Estimated hydraulic conductivities of field-grown Blechnum and Salpichlaena demonstrate that larger conductivities are associated with larger plants. This study contributes toward our knowledge of fern water relations and extends previous growth form/hydraulic architecture characterizations by providing a more comprehensive comparison of closely related species. In addition, this study provides evidence for the relative importance of tracheid characteristics in determining the magnitude of organ hydraulic conductivity.     

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems

Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (K _leaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday K _leaf values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of K _leaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

Hydraulic conductivity of tonoplast-freeChara cells

Summary This study is the first trial to measure the osmotic water permeability or the hydraulic conductivity of the plasmalemma alone of a plant cell. For this purpose tonoplast-free cells were prepared from intenodal cells ofChara australis and their hydraulic conductivities were measured by the transcellular osmosis method.The transcellular hydraulic conductivity did not change after removing the tonoplast. The transcellular hydraulic conductivity of the tonoplast-free cells was dependent on the internal osmotic pressure as is the case in the tonoplast-containing normal cells. The hydraulic conductivities for both endosmosis and exosmosis of the tonoplast-free cells were equal to respective values of the normal cells. Consequently the ratio between the inward and outward hydraulic conductivities did not change due to the loss of the tonoplast. The results indicate that the resistance of the tonoplast to water flow is negligibly small as compared with that of the plasmalemma and further that the tonoplast is not a factor responsible for the direction-dependency of hydraulic conductivity. The hydraulic conductivity of the plasmalemma is invariable for wide variations of K⁺ and Ca²⁺ in the cytoplasm.     

Water uptake by growing cells: an assessment of the controlling roles of wall relaxation, solute uptake, and hydraulic conductance

Growing plant cells increase in volume principally by water uptake into the vacuole. There are only three general mechanisms by which a cell can modulate the process of water uptake: (a) by relaxing wall stress to reduce cell turgor pressure (thereby reducing cell water potential), (b) by modifying the solute content of the cell or its surroundings (likewise affecting water potential), and (c) by changing the hydraulic conductance of the water uptake pathway (this works only for cells remote from water potential equilibrium). Recent studies supporting each of these potential mechanisms are reviewed and critically assessed. The importance of solute uptake and hydraulic conductance is advocated by some recent studies, but the evidence is indirect and conclusions remain controversial. For most growing plant cells with substantial turgor pressure, it appears that reduction in cell turgor pressure, as a consequence of wall relaxation, serves as the major initiator and control point for plant cell enlargement. Two views of wall relaxation as a viscoelastic or a chemorheological process are compared and distinguished.     

Mathematical treatment of water movement in plant cells and tissue: a review

Abstract. Over the past three decades, many contributions have been made to the development of a mathematical basis for describing water transport in plant cells and tissue. This review paper attempts to summarize the more significant contributions and to outline the concepts upon which the various mathematical analyses are founded.
The paper itself is divided into three major sections. Section I deals with the quantitative water relations of single plant cells. Basic equations are developed which describe the water statics and water dynamics of such cells. Included is a discussion of the theory and methods for measuring the various parameters (permeabilities, cell wall elastic moduli, etc.) which enter into the development. The section closes with a presentation of circuit analog models for single plant cells.
Section II is devoted to a review and development of the water relations of plant tissues which contain numerous cells in series. Following a historical overview, various existing models are derived and physical tissue properties which enter the derivation are identified. The concept of 'local equilibrium' is discussed and circuit analog models for single cells are generalized and applied to several cells in series.
The final section contains two example applications of water transport theory as it applies to plant tissue. One application involves radial water movement in a soybean hypocotyl while the other deals with water transport in a growing root tip. A summary at the end of the section is largely devoted to a discussion of the limitations of mathematical models dial are presently available.     

Heterogeneity of soil and soil solution chemistry under Norway spruce (Picea abies Karst.) and European beech (Fagus silvatica L.) as influenced by distance from the stem basis

The present study aims at characterizing plant water status under field conditions on a daily basis, in order to improve operational predictions of plant water stress. Ohm's law analog serves as a basis for establishing daily soil-plant relationships, using experimental data from a water-limited soybean crop: 227-1. The daily transpiration flux, T, is estimated from experimental evapotranspiration data and simulated soil evaporation values. The difference, 227-2, named the effective potential gradient, is derived from i) the midday leaf potential of the uppermost expanded leaves and ii) an effective soil potential accounting for soil potential profile and an effectiveness factor of roots competing for water uptake. This factor is experimentally estimated from field observation of roots. G is an apparent hydraulic conductance of water flow from the soil to the leaves. The value of the lower potential limit for water extraction, required to assess the effective soil potential, is calculated with respect to the plant using the predawn leaf potential. It is found to be equal to –1.2 MPa. It appears that over the range of soil and climatic conditions experienced, the daily effective potential gradient remains constant (1.2 MPa), implying that, on a daily basis, transpiration only depends on the hydraulic conductance. The authors explain this behaviour by diurnal variation of osmotic potential, relying on Morgan's theory (1984). Possible generalization of the results to other crop species is suggested, providing a framework for reasoning plant water behaviour at a daily time step.     

Hydraulic regulation strategies for whole-plant water balance of two maize inbred lines differing in drought resistance under short-term osmotic stress

The conservation of water in agriculture requires an understanding of the mechanisms of plant–water relations. This study aimed to reveal hydraulic regulation strategies of maize (Zea mays L.) for maintaining the plant water balance during drought. The water relations of two maize inbred lines (Tian4 and 478) that differ in their resistance to drought in the field were investigated under well-watered conditions and osmotic stress induced with 10 % PEG 6000. The leaf transpiration rate and leaf water potential of 478 varied diurnally, but remained constant in Tian4, which is more drought resistant. Tian4 plants showed morphological, anatomical and physiological advantages that protected them from foliar water loss. The strategies of leaf hydraulics to regulate leaf water balance during the day and during short-term osmotic stress also differed between Tian4 and 478. The leaf hydraulic conductivity of Tian4 and 478 increased temporarily, but their root hydraulic conductivities were reduced under osmotic stress. However, the root hydraulic conductivity of Tian4 subsequently recovered. Lower and rapidly reduced leaf transpiration and the ability of root hydraulics to recover from short-term osmotic stress can help explain the strategies for plant water balance of drought-tolerant maize.     

不同冲洗液对毛白杨和油松枝条水力导度和抵抗空穴化能力测定值的影响

植物通过木质部管道系统进行水分运输, 木质部的水分运输效率和抗空穴化能力等水力结构特征对于植物物种的分布、抗逆能力等方面起关键性作用。目前, 国内外学者一般采用“冲洗法”进行木质部水力结构研究, 然而在该方法中使用的不同冲洗溶质可能对植物木质部水力结构等产生较大影响, 因此该文研究了3种溶质的冲洗溶液对毛白杨(Populus tomentosa)和油松(Pinus tabulaeformis)枝条的水力导度和抵抗空穴化能力的影响。实验结果表明: 相对于去离子水, 用0.01 mol·L^-1的草酸和0.03 mol·L^-1KCl溶液作为冲洗溶液, 均导致毛白杨木质部导管和油松管胞的水力导度测定值的增大。KCl导致毛白杨和油松木质部抵抗空穴化能力测定值的提高, 草酸导致杨树抵抗空穴化能力测定值增强, 但导致油松抗空穴化能力显著(p<0.01)减弱。小枝水平上, 毛白杨和油松的水分运输效率和抗空穴化能力之间没有显著相关性。另外, 在截枝实验中发现, 毛白杨小枝木质部水力导度随长度增加变化不大, 而油松枝条的木质部水力导度有逐渐增大的趋势。以上的实验结果表明不同溶质下毛白杨和油松枝条的木质部水力导度和抵抗空穴化能力不同, 草酸和KCl可能对木质部管道系统及纹孔处的果胶等产生作用, 从而使毛白杨和油松的水力结构发生变化。毛白杨与油松水力结构在去离子水、草酸和KCl的作用下的不同结果及两物种截枝试验下水力导度的不同变化趋势表明, 导管运输系统和管胞运输系统可能具有不同的水分运输影响因素。     

Nitrate control of root hydraulic properties in plants: translating local information to whole plant response

The sessile lifestyle of plants constrains their ability to acquire mobile nutrients such as nitrate. Whereas proliferation of roots might help in the longer term, nitrate-rich patches can shift rapidly with mass flow of water in the soil. A mechanism that allows roots to follow and capture this source of mobile nitrogen would be highly desirable. Here, we report that variation in nitrate concentration around roots induces an immediate alteration of root hydraulic properties such that water is preferentially absorbed from the nitrate-rich patch. Further, we show that this coupling between nitrate availability and water acquisition results from changes in cell membrane hydraulic properties and is directly related to intracellular nitrate concentrations. Split-root experiments in which nitrate was applied to a portion of the root system showed that the response is both localized and reversible, resulting in rapid changes in water uptake to the portions of the roots exposed to the nitrate-rich patch. At the same time, water uptake by roots not supplied with nitrate was reduced. We believe that the increase in root hydraulic conductance in one part causes a decline of water uptake in the other part due to a collapse in the water potential gradient driving uptake. The translation of local information, in this case nitrate concentration, into a hydraulic signal that can be transmitted rapidly throughout the plant and thus coordinate responses at the whole plant level, represents an unexpected, higher level physiological interaction that precedes the level of gene expression.     

An exceptional role for flowering plant physiology in the expansion of tropical rainforests and biodiversity

Movement of water from soil to atmosphere by plant transpiration can feed precipitation, but is limited by the hydraulic capacities of plants, which have not been uniform through time. The flowering plants that dominate modern vegetation possess transpiration capacities that are dramatically higher than any other plants, living or extinct. Transpiration operates at the level of the leaf, however, and how the impact of this physiological revolution scales up to the landscape and larger environment remains unclear. Here, climate modelling demonstrates that angiosperms help ensure aseasonally high levels of precipitation in the modern tropics. Most strikingly, replacement of angiosperm with non-angiosperm vegetation would result in a hotter, drier and more seasonal Amazon basin, decreasing the overall area of ever-wet rainforest by 80 per cent. Thus, flowering plant ecological dominance has strongly altered climate and the global hydrological cycle. Because tropical biodiversity is closely tied to precipitation and rainforest area, angiosperm climate modification may have promoted diversification of the angiosperms themselves, as well as radiations of diverse vertebrate and invertebrate animal lineages and of epiphytic plants. Their exceptional potential for environmental modification may have contributed to divergent responses to similar climates and global perturbations, like mass extinctions, before and after angiosperm evolution.     

The units of currency for plant water status

Abstract. The appropriate measure for plant water status is of fundamental importance in plant water relations. One possible approach is to consider a hypothetical sensor for water status at the cell level. It is argued that such a sensor must respond to an intrinsic variable describing the quantity of water present in the cell. Uncertainty about the variable being sensed has affected our interpretion of plant responses to drought. In general, a more rigorous analysis of the effects of tissue hydraulic parameters and further understanding of the pertinent variable for water status are needed to assess such questions. For example, the significance of changes in the elasticity of cell walls in response to drought depends on the measure for water status. A simple model for water transport suggests that a decrease in the elastic modulus may act to maintain turgor for succulent plant species, but it is not clear how increases in the elastic modulus enhance water uptake for non-succulent species.     

The differential role of mechanisms for drought resistance in a Mediterranean evergreen shrub: a simulation approach

Plants have the ability to dampen the effects of variability in water resources. Various mechanisms contribute to these properties: reduction of leaf area, increased rooting depth and stomatal conductance. To evaluate the differential roles and interactions of these mechanisms, we have built a model and simulated flows of water in Mediterranean evergreen scrub. The essential concept of this model is that the water status of the canopy is governed by the water lost by transpiration, the availability of soil water and the hydraulic resistances to water flow in soil and plant. The amount of water supplied by the roots is related to changes in water potential between the soil and the leaf. The amount of water lost to the atmosphere is regulated by an interaction between atmospheric demand and canopy water potential. Water uptake by plant is assumed equal to plant water loss. Leaf area appears to affect largely the annual water balance. The critical leaf water potential required to reduce the maximum stomatal conductance by half has a dominant effect on annual leaf water potential. Reducing rooting depth induces a new functional equilibrium for the plant. This new equilibrium is reached by decreasing leaf area and the critical leaf water potential. Our results show the complexity of interactions of these mechanisms and highlight the importance of the coordination between them. Finally, we suggest a reconsideration of these mechanisms in a context of the survival and long-term persistence of the plant.     

干旱胁迫植物个体生理响应及其生态模型预测研究进展

全球气候突变导致干旱事件频发,进而易引发严重的植物衰退甚至死亡,聚焦植物尤其是树木死亡的生理学机制并期望基于此评估及预测气候变化导致植物死亡风险已成为热点话题。植物通过调整内在生理代谢过程,例如通过调节渗透物质的含量,来平衡渗透势、维持细胞膨压、调节植物激素的信号水平,诱导植物气孔开放程度降低,有利于植物保存水分、调控植物水通道蛋白的表达,进而保持体内水分稳定并对干旱胁迫做出快速响应。这些生理过程中的每一环调节都为了确保水分运输的效率和安全性,增加植物抗旱性以及生态系统稳定性。植物的抗旱性不仅体现在生理代谢方面的调节,还表现在植物水力特性与解剖结构间相辅相成。当植物改变水力特性的同时,其茎叶会在解剖结构上做出调整以满足植物在干旱环境下水分供需平衡,从而降低植物蒸腾水分散失、增强细胞储水并提高生存能力。植物应对水分胁迫的策略通常与水分消耗和碳获取之间的平衡有关,明晰植物水分消耗与光合碳获取间存在平衡关系的性状特征便于更好地理解植物的水分利用策略。然而,植物表现出的任意单一性状特征的强弱都无法代表整个植物适应逆境的优劣,未来只有通过将植物更多性状特征进行相互关联,以具有代表植物水力功能、结...     

Ascorbate peroxidase activity, not the mRNA level, is enhanced in salt-stressed Raphanus sativus plants

Important functions of water relations are considered to be related to genome size diversity in gymnosperms. We investigated relationships among genome size, dimensional characteristics of conductive cells, and water relations parameters by using young seedlings of six Pinus species. Xylem hydraulic conductivity was not correlated with genome size and dimensional characteristics of conductive cells, but the water potential at the turgor loss point was. Pinus species with large genome sizes had thick cell walls and small ratios of lumen radii to cell wall thickness in their conductive cells, and those species lost their turgor to tissue dehydration al low water potentials. The characteristics observed in the present study may contribute to pine drought tolerance.     

Hydraulic Conductivity Recovery versus Water Pressure in Xylem of Acer saccharum

Experiments were conducted to determine the influence of stem diameter, xylem pressure potential, and temperature on the rate of recovery of hydraulic conductivity in embolized stems of Acer saccharum Marsh. Recovery of conductivity was accompanied by an increase in stem water content as water replaced air bubbles and bubbles dissolved from vessels into the surrounding water. The time required for stems to go from less than 3 to 100% hydraulic conductivity increased approximately with the square of the stem diameter and increased with decreasing xylem pressure potential. Recovery was halted when xylem pressure potential decreased below −6 kPa. Increasing xylem pressure from 13 to 150 kPa reduced the time for recovery by a factor of 4. Temperature had little influence on the rate of recovery of hydraulic conductivity. All of these results are in accord with a theory of bubble dissolution in which it is assumed that: (a) the rate of bubble dissolution is rate limited by diffusion of air from the bubbles to the outer surface of the stems, (b) the equilibrium concentration of gases in liquid in stems is determined by Henry's law at all air-water interfaces, (c) the equilibrium solubility concentration is determined only by the partial pressure of the gas in the gas phase and not directly by the liquid-phase pressure, and (d) the gas pressure of an entrapped air bubble in the lumen of a cell can never be less than atmospheric pressure at equilibrium.     

Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park,Zhejiang Province,China

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.