Relationships among xylem transport, biomechanics and storage in stems and roots of nine Rhamnaceae species of the California chaparral

Here, hypotheses about stem and root xylem structure and function were assessed by analyzing xylem in nine chaparral Rhamnaceae species. Traits characterizing xylem transport efficiency and safety, mechanical strength and storage were analyzed using linear regression, principal components analysis and phylogenetic independent contrasts (PICs). Stems showed a strong, positive correlation between xylem mechanical strength (xylem density and modulus of rupture) and xylem transport safety (resistance to cavitation and estimated vessel implosion resistance), and this was supported by PICs. Like stems, greater root cavitation resistance was correlated with greater vessel implosion resistance; however, unlike stems, root cavitation resistance was not correlated with xylem density and modulus of rupture. Also different from stems, roots displayed a trade-off between xylem transport safety from cavitation and xylem transport efficiency. Both stems and roots showed a trade-off between xylem transport safety and xylem storage of water and nutrients, respectively. Stems and roots differ in xylem structural and functional relationships, associated with differences in their local environment (air vs soil) and their primary functions.     

Water uptake profile response of corn to soil moisture depletion

The effects of soil moisture distribution on water uptake of drip‐irrigated corn were investigated by simultaneously monitoring the diurnal evolution of sap flow rate in stems, of leaf water potential, and of soil moisture, during intervals between successive irrigations. The results invalidate the steady‐state resistive flow model for the continuum. High hydraulic capacitance of wet soil and low hydraulic conductivity of dry soil surrounding the roots damped significantly diurnal fluctuations of water flow from bulk soil to root surface. By contrast, sap flow responded directly to the large diurnal variation of leaf water potential. In wet soil, the relation between the diurnal courses of uptake rates and leaf water potential was linear. Water potential at the root surface remained nearly constant and uniformly distributed. The slope of the lines allowed calculating the resistance of the hydraulic path in the plant. Resistances increased in inverse relation with root length density. Soil desiccation induced a diurnal variation of water potential at the root surface, the minimum occurring in the late afternoon. The increase of root surface water potential with depth was directly linked to the soil desiccation profile. The development of a water potential gradient at the root surface implies the presence of a significant axial resistance in the root hydraulic path that explains why the desiccation of the soil upper layer induces an absolute increase of water uptake rates from the deeper wet layers.     

Limitation of plant water use by rhizosphere and xylem conductance: results from a model

Hydraulic conductivity ( K ) in the soil and xylem declines as water potential ( Ψ ) declines. This results in a maximum rate of steady-state transpiration ( E _crit) and corresponding minimum leaf Ψ ( Ψ _crit) at which K has approached zero somewhere in the soil–leaf continuum. Exceeding these limits causes water transport to cease. A model determined whether the point of hydraulic failure (where K = 0) occurred in the rhizosphere or xylem components of the continuum. Below a threshold of root:leaf area ( A _R: A _L), the loss of rhizosphere K limited E _crit and Ψ _crit. Above the threshold, loss of xylem K from cavitation was limiting. The A _R: A _L threshold ranged from > 40 for coarse soils and/or cavitation-resistant xylem to < 0·20 in fine soils and/or cavitation-susceptible xylem. Comparison of model results with drought experiments in sunflower and water birch indicated that stomatal regulation of E reflected the species' hydraulic potential for extracting soil water, and that the more sensitive stomatal response of water birch to drought was necessary to avoid hydraulic failure. The results suggest that plants should be xylem-limited and near their A _R: A _L threshold. Corollary predictions are (1) within a soil type the A _R: A _L should increase with increasing cavitation resistance and drought tolerance, and (2) across soil types from fine to coarse the A _R: A _L should increase and maximum cavitation resistance should decrease.     

Resistance to root growth and changes in the concentrations of ABA within the root and xylem sap during root-restriction stress

The possibility that increased soil resistance to root growth may mediate the dwarfing response associated with root-restriction stress (RRS), via an abscisic acid (ABA) transduction mechanism, was investigated by characterizing the responses of tomato plants (Lycopersicon esculentum Mill cv. Red Dwarf) and changes within the soil environment at three rooting volumes (RV) (200, 400 and 800 cm³). Plant dry weight, leaf area and stomatal conductance decreased with RRS, although leaf water potential was unaffected by RRS. The concentration of ABA within the root system ([ABA]rt) and xylem sap ([ABA]xy) increased with RRS. Increased bulk density caused soil resistance to root growth to increase with increasing RRS. Changes in the soil environment, other than bulk density, which may have induced this variation in concentrations of ABA, were either eliminated or shown not to limit plant growth. The proportional relationships between RRS and soil resistance, [ABA]rt and [ABA]xy, and the inverse relationship between RRS and plant growth, are possibly indicative of the restricted root system experiencing increased resistance to root growth, with the subsequent initiation of a cascade of growth inhibiting responses.     

树木高生长限制的几个假说

树木生长到一定年龄后高生长停滞,对这一现象的解释存在很多争议.成熟假说认为树木顶端分生组织分裂活性下降导致树木高生长减慢.营养限制假说认为土壤中营养元素(特别是氮素)在植物活体或枯落物中积累使土壤中可利用的养分含量降低,细根生物量增加和叶片光合能力下降导致了地上部分生长的减缓.呼吸假说认为边材呼吸消耗随个体发育的增加使投入到高生长的碳减少.水力限制假说认为水分运输阻力随高度增加而增加导致了叶片总光合碳同化下降,分配到高生长的生物量减少.树木发展假说认为植物用多种调节机制克服随个体发育增加的水力阻力,包括叶片结构和生理特征的变化,叶/边材面积比降低,边材渗透性和树干储水能力的增加等.水力限制假说得到了较多的关注,对不同高度树木的叶比导率、光合特征和树干生长量等测定结果支持这一假说.但对这一假说也存在很多的争议,主要表现在:水力阻力是否确实随高度的增加而增加,水力阻力的分布,补偿机制的作用和生物量分配转变等.本文综述了树木高生长限制的4个假说以及争论的焦点,并总结了目前研究的热点问题和今后的研究方向.     

树木高生长限制的几个假说

树木生长到一定年龄后高生长停滞, 对这一现象的解释存在很多争议。成熟假说认为树木顶端分生组织分裂活性下降导致树木高生长减慢。营养限制假说认为土壤中营养元素(特别是氮素)在植物活体或枯落物中积累使土壤中可利用的养分含量降低, 细根生物量增加和叶片光合能力下降导致了地上部分生长的减缓。呼吸假说认为边材呼吸消耗随个体发育的增加使投入到高生长的碳减少。水力限制假说认为水分运输阻力随高度增加而增加导致了叶片总光合碳同化下降, 分配到高生长的生物量减少。树木发展假说认为植物用多种调节机制克服随个体发育增加的水力阻力, 包括叶片结构和生理特征的变化, 叶/边材面积比降低, 边材渗透性和树干储水能力的增加等。水力限制假说得到了较多的关注, 对不同高度树木的叶比导率、光合特征和树干生长量等测定结果支持这一假说。但对这一假说 也存在很多的争议, 主要表现在: 水力阻力是否确实随高度的增加而增加, 水力阻力的分布, 补偿机制的作用和生物量分配转变等。本文综述了树木高生长限制的4个假说以及争论的焦点, 并总结了目前研究的热点问题和今后的研究方向。     

Microtensiometer technique for in situ measurement of soil matric potential and root water extraction from a sandy soil

The suitability of microtensiometers to measure the spatial variation of soil matric potential and its diurnal change was tested in a pot experiment with pearl millet (Pennisetum americanum [L.] Leeke) in a sandy soil as the soil dried out.The temporal and spatial resolution of this technique allowed precise measurement of soil matric potential and thus estimation of soil water extraction from different compartments as well as from the whole rooting zone. The technique also allowed the measurement of rehydration of plants at night and root water uptake rate per unit soil volume or per unit root length. The precision of determination of root water uptake depended greatly on the accuracy of the estimate of hydraulic conductivity, which was derived from a bare soil and might be different for a cropped soil owing to aggregation induced by the root system. A linear relationship between root length and water uptake was found (r²=0.82), irrespective of variation in soil water content between compartments and despite the variation in root age, xylem differentiation and suberin formation expected to exist between different compartments of the rooting zone. As the experiment was carried out in a range of soil matric potentials between –4 and –30 kPa, drought stress did not occur. Further information at lower soil matric potentials are required, to address questions such as the importance of soil resistance for water uptake, or which portion of the root system has to be stressed to induce hormonal signals to the shoot. The microtensiometer technique can be applied to soil matric potentials up to –80 kPa.     

The water relations of the root–soil interface

The difference in hydrostatic pressure between the xylem of the leaf and the soil depends, for a given transpiration rate, on the series of hydraulic resistances encountered along this pathway. Many studies have shown that the sum of the resistances in the plant and the soil is too small to account for the fall in water pressure between the leaf xylem and the soil, especially when plants are growing in sandy soils, which are prone to dry rapidly. A resistance at the root–soil interface, caused possibly by poor contact between the roots and the soil, has been proposed to account for the discrepancy. We explored the resistance in the pathway from soil to leaf using a technique that allows precise and continuous non-destructive measurement of the hydrostatic pressure in the leaf xylem. When the soil was leached with water, the fall in leaf water status as the soil dried was reasonably well described by a simple physical model without the need to invoke an interfacial resistance. However, when the soil was flushed with a nutrient solution with an osmotic pressure of 70kPa, the hydrostatic pressure in the leaf xylem fell several times faster than that in the soil. We suggest that solutes accumulated either in the root or just outside it, creating large osmotic pressures, which gave the appearance of an interfacial resistance.     

Electrical resistances of corn root segments

Longitudinal electrical resistances have been measured on 2-centimeter segments of corn (Zea mays L.) roots, cut at varying distances from the root apex. The segment resistances vary from 400 to 100 kilohms per centimeter along the root length (apex to 18 cm), with the maximum occurring in the 2- to 4-centimeter segment, and decreasing thereafter toward the root base. Measurements of isolated root cortical sleeves and steles show that the pathway of least resistance is in the cortex, which has a greater cross-sectional area; the specific resistance of the older stele is less than that of the cortex. The anatomical state of the xylem cannot be inferred from electrical resistance determinations.     

Oscillations of electrical potential along a root of a higher plant

Higher plants exhibit an oscillation of electrical potential near the surface along the root. The oscillation was studied with the aid of both the usual intracellular microelectrode technique and the extracellular multielectrode technique, the latter making it possible to measure simultaneously electrical potentials along the root. It was found that the oscillation of extracellular surface potential showed the largest amplitude in the elongation region, and the phase of the oscillation in this region differed by 180 degrees from that in the mature region, where the oscillation appeared coherent. The measurement of the intracellular electrical potential suggested the existence of oscillatory components localized to the parenchyma/xylem interface in the elongation region. A theoretical analysis based on an electrical circuit network described the above-mentioned behavior. It was shown that the oscillation was propagated along the root over several centimeters without substantial decay in the mature region.     

Leaf water stress in engelmann spruce: influence of the root and shoot environments

The response of xylem pressure potential of Engelmann spruce (Picea engelmannii Engelm.) to environmental factors was studied in the natural subalpine environment. Data were analyzed in the context of a leaf water potential model based upon the van den Honert model for water transport through the soil-plant-atmosphere continuum. At soil temperatures of 10 to 15 C, xylem pressure potential decreased to about −10 bars as the ratio of leaf to air absolute humidity difference to leaf diffusion resistance (an estimate of transpiration) increased. The potentials were slightly lower at all flux rates above zero when the soil temperature was 5 to 10 C, and at temperatures of 0 to 5 C the potentials decreased sharply to as low as −20.4 bars, even though the soil water supply was adequate. The relative viscosity of water and soil to leaf resistances for flow were compared for Engelmann spruce and citrus at low soil temperatures. These comparisons indicated that decreased root permeability was probably not an important factor causing higher stresses in spruce at 5 to 10 C, but for citrus, root permeability became limiting at soil temperatures as high as 13.5 C. Xylem pressure potential was correlated with net radiation during the daytime when soil temperature was above 7 C. Under other conditions, however, xylem potential and net radiation apparently had a different relationship. The relationship between flux density and potential was the same on unshaded and shaded portions of the crown, with differences in potential related to differences in flux density.     

Modelling non-uniform soil water uptake by a single plant root

The assumption of uniform water flow to the root or uniform water potential at the root surface was shown by Hainsworth and Aylmore (1986, 1989) to be erroneous. The present paper demonstrates how the non-uniform uptake of water by a single root can be modeled. Differential equations are numerically solved to describe simultaneous water movement in the plant and in the soil. In the plant, boundary conditions are the water potentials at the root surface (Ψ_s) and in the xylem at the root base (Ψ_b). A set of difference equations describe the flow of water radially through the cortex to the xylem and in the xylem axially upwards to the base. For calculating the water flow in the soil and the values of Ψ_s, i.e. the boundary conditions for flow in the root, the finite element method (FEM) is used, the boundary conditions being the flux of water into the plant root and the zero flow across the wall, bottom and surface of a hypothetical soil cylinder surrounding the root. ei]Section editor: B E Clothier     

Root-shoot interactions in mineral nutrition

In this paper four classes of co-operative root-shoot interations are addressed. (I) Nitrogen concentrations in the xylem sap originating from the root and in the phloem sap as exported from source leaves are much lower than those required for growth by apices and developing organs. Enrichment of xylem sap N is achieved by xylem to xylem (X-X) transfer, by which reduced N, but not nitrate, is abstracted from the xylem of leaf traces and loaded into xylem vessels serving the shoot apex. Nitrogen enrichment of phloem sap from source leaves is enacted by transfer of reduced N from xylem to phloem (X-P transfer). Quantitative data for the extent of the contribution of X-X and X-P transfer to the nutrition of young organs of Ricinus communis L. and for their change with time are presented. (II) Shoot and root cooperate in nitrate reduction and assimilation. The partitioning of this process between shoot and root is shifted towards the root under conditions of nitrate- and K-deficiency and under salt stress, while P deficiency shifts nitrate reduction almost totally to the shoot. All four changes in partitioning can be attributed to the need for cation-anion balance during xylem transport and the change in electrical charge occurring with nitrate reduction. (III) Even maintenance of the specificity of ion uptake by the root may – in addition to its need for energy – require a shoot-root interaction. This is shown to be needed in the case of the maintenance of K/Na selectivity under the highly adverse condition of salt stress and absence of K supply from the soil. (IV) Hormonal root to shoot interactions are required in the whole plant for sensing mineral imbalances in the soil. This is shown and addressed for conditions of salt stress and of P deficiency, both of which lead to a strong ABA signalling from root to shoot but result in different patterns of response in the shoot.     

Response of liquid flow resistance to soil drying in seedlings of four deciduous angiosperms

Summary Soil-leaf resistance to liquid water flow (R) in moist and drying soil was compared in three-month-old seedlings of two drought tolerant (white [Quercus alba L.], post oak [Q. stellata Wangenh.]) and two drought sensitive forest species (sugar maple [Acer saccharum Marsh.], black walnut [Juglans nigra L.]). At high soil moisture (_s–0.3 MPa), R was higher in J. nigra than in the other species, and as soil water was depleted R increased most in this species. In contrast, the lowest resistance at all levels of soil moisture was observed in Q. stellata. At _s of –1.5 MPa, R of drought-sensitive J. nigra and A. saccharum was about twice as high as that of the two drought-tolerant Quercus species. The difference in R between the two Quercus species was much smaller than that between this pair and the other two species. These differences among species in flow resistance may be attributable to: 1) variation in the balance between root surface area and leaf area, 2) variation in the inherent absorption capacity of the root systems and in xylem water conducting systems or 3) differences in root permeability, shrinkage and mortality in severely stressed seedlings. As the soil dried, seedlings of all species exhibited pronounced reductions in transpiration rate, which prevented development of large water potential gradients between leaves and the soil. Reduction in transpiration in J. nigra was especially pronounced, resulting in a decrease in the soil-to-leaf water potential gradient in dry soil despite high flow resistance. The observed differences among species in flow resistance are correlated with natural distribution patterns.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Analysis of Root Growth by Impedance Spectroscopy (EIS)

Electrical impedance spectroscopy (EIS) is investigated as a non-destructive method for monitoring root growth of tomato. This paper aims to (i) review the basic principles of EIS applied to the characterisation of the different parts of the soil–root–stem-electrode continuum, (ii) experiment the validity of the relationship between root weight and root capacitance taking into account the influence of the soil and plant electrodes position, (iii) describe an EIS analysis of the root growth of tomato plants. All experiments were carried out in 50 dm³ containers either in hydroponics at 930 μS for the test of root fresh or dry weight and root capacitance relationships, or in a potting mix (oxisol) for electrode placement tests and EIS estimation of root growth. Electrical measurements of the soil–root–stem-electrode continuum were done with a two-electrode measuring system using unpolarisable Ag–AgCl electrodes. A ‘root cutting’ and a ‘progressively immersed root system’ experiments were carried out in order to validate the relationship between root capacitance and root mass at 1 kHz. The effects of soil electrode and plant electrode placement were also tested, pointing out the sensitivity of the method to the insertion height of the “plant electrode” into the stem. For the root growth experiment, Impedance Spectra (IS) measurements were made just before harvesting the roots for dry weight and length determination. Measurements were made 14, 22, 26 and 39 days after planting, until flowering. The IS of the soil–root–stem-electrode continuum was modelled by a lumped electric circuit consisting of a series resistor R ₀ for the soil and of four parallel resistance (R _i )-capacitance (C _i ) circuits for the other components of the circuit. The model had nine parameters whose values were estimated by Complex Nonlinear Least Squares curve fitting. A stepwise ascendant regression was used to identify the electrical parameters that better correlated with root dry mass or length increment: C ₃ and C ₄ were well correlated with root dry mass with a r ² of 0.975, whereas root length was explained by the combination of 1/R ₃, C ₃, 1/R ₂ and 1/R ₁ with a r ² of 0.986. This work may be considered as a new methodological contribution to the understanding of root electrical properties in the non-destructive diagnosis of root systems.     

The influence of slope on <Emphasis Type="Italic">Spartium junceum</Emphasis> root system: morphological,anatomical and biomechanical adaptation

Root systems have a pivotal role in plant anchorage and their mechanical interactions with the soil may contribute to soil reinforcement and stabilization of slide-prone slopes. In order to understand the responses of root system to mechanical stress induced by slope, samples of Spartium junceum L., growing in slope and in plane natural conditions, were compared in their morphology, biomechanical properties and anatomical features. Soils sampled in slope and plane revealed similar characteristics, with the exception of organic matter content and penetrometer resistance, both higher in slope. Slope significantly influenced root morphology and in particular the distribution of lateral roots along the soil depth. Indeed, first-order lateral roots of plants growing on slope condition showed an asymmetric distribution between up- and down-slope. Contrarily, this asymmetric distribution was not observed in plants growing in plane. The tensile strength was higher in lateral roots growing up-slope and in plane conditions than in those growing down-slope. Anatomical investigations revealed that, while roots grown up-slope had higher area covered by xylem fibers, the ratio of xylem and phloem fibers to root diameter did not differ among the three conditions, as also, no differences were found for xylem fiber cell wall thickness. Roots growing up-slope were the main contributors to anchorage properties, which included higher strength and higher number of fibers in the xylematic tissues. Results suggested that a combination of root-specific morphological, anatomical and biomechanical traits, determines anchorage functions in slope conditions.     

An Electrical Analogue to Simulate the Oxygen Relations of Roots in Anaerobic Media

The paper describes the construction and the theory underlying a simple electrical analogue which may be used to simulate the internal oxygen relations of roots in the wetland condition. The model represents an unbranched root lying within an oxygen sink and consists of twenty circuit units linked in series. Each unit which can be programmed independently, represents the major oxygen diffusion paths, impedances and sinks recognizable in a centimetre-long segment of root/wet soil system. Simplicity has been achieved by specifying constant radius for the root (0.05 cm), constant potential activity for the external sink, by expressing respiration and radial oxygen loss as a function of root surface area, and by the non-inclusion of oxygen storage capacity.     

Drought resistance in early and late secondary successional species from a tropical dry forest: the interplay between xylem resistance to embolism,sapwood water storage and leaf shedding

The mechanisms of drought resistance that allow plants to successfully establish at different stages of secondary succession in tropical dry forests are not well understood. We characterized mechanisms of drought resistance in early and late‐successional species and tested whether risk of drought differs across sites at different successional stages, and whether early and late‐successional species differ in resistance to experimentally imposed soil drought. The microenvironment in early successional sites was warmer and drier than in mature forest. Nevertheless, successional groups did not differ in resistance to soil drought. Late‐successional species resisted drought through two independent mechanisms: high resistance of xylem to embolism, or reliance on high stem water storage capacity. High sapwood water reserves delayed the effects of soil drying by transiently decoupling plant and soil water status. Resistance to soil drought resulted from the interplay between variations in xylem vulnerability to embolism, reliance on sapwood water reserves and leaf area reduction, leading to a tradeoff of avoidance against tolerance of soil drought, along which successional groups were not differentiated. Overall, our data suggest that ranking species' performance under soil drought based solely on xylem resistance to embolism may be misleading, especially for species with high sapwood water storage capacity.     

Biomass and morphology of fine roots in temperate broad-leaved forests differing in tree species diversity: is there evidence of below-ground overyielding?

Biodiversity effects on ecosystem functioning in forests have only recently attracted increasing attention. The vast majority of studies in forests have focused on above-ground responses to differences in tree species diversity, while systematic analyses of the effects of biodiversity on root systems are virtually non-existent. By investigating the fine root systems in 12 temperate deciduous forest stands in Central Europe, we tested the hypotheses that (1) stand fine root biomass increases with tree diversity, and (2) ‘below-ground overyielding’ of species-rich stands in terms of fine root biomass is the consequence of spatial niche segregation of the roots of different species. The selected stands represent a gradient in tree species diversity on similar bedrock from almost pure beech forests to medium-diverse forests built by beech, ash, and lime, and highly-diverse stands dominated by beech, ash, lime, maple, and hornbeam. We investigated fine root biomass and necromass at 24 profiles per stand and analyzed species differences in fine root morphology by microscopic analysis. Fine root biomass ranged from 440 to 480 g m⁻² in the species-poor to species-rich stands, with 63–77% being concentrated in the upper 20 cm of the soil. In contradiction to our two hypotheses, the differences in tree species diversity affected neither stand fine root biomass nor vertical root distribution patterns. Fine root morphology showed marked distinctions between species, but these root morphological differences did not lead to significant differences in fine root surface area or root tip number on a stand area basis. Moreover, differences in species composition of the stands did not alter fine root morphology of the species. We conclude that ‘below-ground overyielding’ in terms of fine root biomass does not occur in the species-rich stands, which is most likely caused by the absence of significant spatial segregation of the root systems of these late-successional species.