Conductivity Of Root System Research Articles

Root System Architecture Reorganization Under Decreasing Soil Phosphorus Lowers Root System Conductance of Zea mays.

The global supply of phosphorus is decreasing. At the same time, climate change reduces the availability of water in most regions of the world. Insights on how decreasing phosphorus availability influences plant architecture are crucial to understanding its influence on plant functional properties, such as the root system's water uptake capacity. In this study, we investigated the structural and functional responses of Zea mays to varying phosphorus fertilization levels focusing especially on the root system's conductance. A rhizotron experiment with soils ranging from severe phosphorus deficiency to sufficiency was conducted. We measured the architectural parameters of the whole plant and combined them with root hydraulic properties to simulate time-dependent root system conductance of growing plants under different phosphorus levels. We observed changes in the root system architecture, characterised by decreasing crown root elongation and reduced axial root radii with declining phosphorus availability. Modeling revealed that only plants with optimal phosphorus availability sustained a high root system conductance, while all other phosphorus levels led to a significantly lower root system conductance, both under light and severe phosphorus deficiency. We postulate that phosphorus deficiency decreases root system conductance, which could mitigate drought conditions through a more conservative water use strategy, but ultimately reduces biomass and impairs root development and overall water uptake capacity. Our results also highlight that the organisation of the root system, rather than its overall size, is critical for estimating important root functions.

Root hydraulic properties: An exploration of their variability across scales.

Root hydraulic properties are key physiological traits that determine the capacity of root systems to take up water, at a specific evaporative demand. They can strongly vary among species, cultivars or even within the same genotype, but a systematic analysis of their variation across plant functional types (PFTs) is still missing. Here, we reviewed published empirical studies on root hydraulic properties at the segment-, individual root-, or root system scale and determined its variability and the main factors contributing to it. This corresponded to a total of 241 published studies, comprising 213 species, including woody and herbaceous vegetation. We observed an extremely large range of variation (of orders of magnitude) in root hydraulic properties, but this was not caused by systematic differences among PFTs. Rather, the (combined) effect of factors such as root system age, driving force used for measurement, or stress treatments shaped the results. We found a significant decrease in root hydraulic properties under stress conditions (drought and aquaporin inhibition, p < .001) and a significant effect of the driving force used for measurement (hydrostatic or osmotic gradients, p < .001). Furthermore, whole root system conductance increased significantly with root system age across several crop species (p < .01), causing very large variation in the data (>2 orders of magnitude). Interestingly, this relationship showed an asymptotic shape, with a steep increase during the first days of growth and a flattening out at later stages of development. We confirmed this dynamic through simulations using a state-of-the-art computational model of water flow in the root system for a variety of crop species, suggesting common patterns across studies and species. These findings provide better understanding of the main causes of root hydraulic properties variations observed across empirical studies. They also open the door to better representation of hydraulic processes across multiple plant functional types and at large scales. All data collected in our analysis has been aggregated into an open access database (https://roothydraulic-properties.shinyapps.io/database/), fostering scientific exchange.

Expansion and evaluation of two coupled root–shoot models in simulating CO2 and H2O fluxes and growth of maize

AbstractLeaf water pressure head (Ψleaf) and more specifically its critical thresholds (Ψthreshold) characterize stomatal control of transpiration, particularly for C4 plants, but this physiological process has rarely been integrated into dynamic crop models at the field scale. We further extended two coupled models with Feddes root water uptake (RWU) and Couvreur RWU models by adding the C4 photosynthesis model in order to be applicable to a maize (Zea mays L.) crop growing under field conditions. The Feddes RWU model relates RWU and plant stress directly to the root zone water hydraulic potential, whereas the Couvreur model explicitly represents hydraulic signals between soil and stomata. Model performance was evaluated with a comprehensive dataset including stomatal conductance, Ψleaf, sap flow, gross assimilation rate (GAR), soil water content (SWC), dry biomass, and leaf area index (LAI) from a two‐season maize experiment under contrasting environments and water regimes. For the Couvreur model, the RWU and dry biomass were more sensitive to the root hydraulic conductance parameters than to Ψthreshold and root growth parameters. The agreement index (I) for the Couvreur model after calibration was 0.91, 0.80, 0.81, and 0.69 for biomass, LAI, RWU, and GAR, respectively. The Feddes model performed similarly for the same metrics. The Feddes model simulated accurately the plant water stress in the first 45 d of the growing season, whereas the Couvreur model inaccurately predicted water stress, which resulted in lower agreement with observations (i.e., RMSEs of biomass simulated by Feddes and Couvreur model averaged from four validated plots were 0.171 and 0.214 kg m–2, respectively). The Feddes model showed the potential to be used for maize under water stress whereas the Couvreur model needs to be further evaluated and improved with an adequate estimation of root hydraulic conductance. A dynamic parameterization of normalized root system conductance and/or more accurate assimilate allocation to the roots, especially under drought stress, should be considered in order to apply this model in future studies to maize.

From hydraulic root architecture models to macroscopic representations of root hydraulics in soil water flow and land surface models

Abstract. Root water uptake is an important process in the terrestrial water cycle. How this process depends on soil water content, root distributions, and root properties is a soil–root hydraulic problem. We compare different approaches to implement root hydraulics in macroscopic soil water flow and land surface models. By upscaling a three-dimensional hydraulic root architecture model, we derived an exact macroscopic root hydraulic model. The macroscopic model uses the following three characteristics: the root system conductance, Krs, the standard uptake fraction, SUF, which represents the uptake from a soil profile with a uniform hydraulic head, and a compensatory matrix that describes the redistribution of water uptake in a non-uniform hydraulic head profile. The two characteristics, Krs and SUF, are sufficient to describe the total uptake as a function of the collar and soil water potential, and water uptake redistribution does not depend on the total uptake or collar water potential. We compared the exact model with two hydraulic root models that make a priori simplifications of the hydraulic root architecture, i.e., the parallel and big root model. The parallel root model uses only two characteristics, Krs and SUF, which can be calculated directly following a bottom-up approach from the 3D hydraulic root architecture. The big root model uses more parameters than the parallel root model, but these parameters cannot be obtained straightforwardly with a bottom-up approach. The big root model was parameterized using a top-down approach, i.e., directly from root segment hydraulic properties, assuming a priori a single big root architecture. This simplification of the hydraulic root architecture led to less accurate descriptions of root water uptake than by the parallel root model. To compute root water uptake in macroscopic soil water flow and land surface models, we recommend the use of the parallel root model with Krs and SUF computed in a bottom-up approach from a known 3D root hydraulic architecture.

MARSHAL, a novel tool for virtual phenotyping of maize root system hydraulic architectures

Abstract Functional-structural root system models combine functional and structural root traits to represent the growth and development of root systems. In general, they are characterized by a large number of growth, architectural and functional root parameters, generating contrasted root systems evolving in a highly non-linear environment (soil, atmosphere), which makes the link between local traits and functioning unclear. On the other end of the root system modelling continuum, macroscopic root system models associate to each root system a set of plant-scale, easily interpretable parameters. However, as of today, it is unclear how these macroscopic parameters relate to root-scale traits and whether the upscaling of local root traits is compatible with macroscopic parameter measurements. The aim of this study was to bridge the gap between these two modelling approaches. We describe here the MAize Root System Hydraulic Architecture soLver (MARSHAL), a new efficient and user-friendly computational tool that couples a root architecture model (CRootBox) with fast and accurate algorithms of water flow through hydraulic architectures and plant-scale parameter calculations. To illustrate the tool’s potential, we generated contrasted maize hydraulic architectures that we compared with root system architectural and hydraulic observations. Observed variability of these traits was well captured by model ensemble runs. We also analysed the multivariate sensitivity of mature root system conductance, mean depth of uptake, root system volume and convex hull to the input parameters to highlight the key model parameters to vary for virtual breeding. It is available as an R package, an RMarkdown pipeline and a web application.

Whole root system water conductance responds to both axial and radial traits and network topology over natural range of trait variation

Current theory and supporting research suggests that radial transport is the most limiting factor to root water uptake, raising the question whether only absorbing root length and radial conductivity matter to water uptake. Here, we extended the porous pipe analytical model of root water uptake to entire root networks in 3D and analysed the relative importance of axial and radial characteristics to total uptake over parameter ranges reported in the literature. We found that network conductance can be more sensitive to axial than radial conductance of absorbing roots. When axial transport limits uptake, more dichotomous topology, especially towards the base of the network, increases water uptake efficiency, while the effect of root length is reduced. Whole root system conductance was sensitive to radial transport and length in model lupin (Lupinus angustifolius L.), but to axial transport and topology in wheat (Triticum aestivum L.), suggesting the root habit niche space of monocots may be constrained by their loss of secondary growth. A deep tap root calibrated to oak (Quercus fusiformis J. Buchholz) hydraulic parameters required 15 times more xylem volume to transport comparable amounts of water once recalibrated to parameters from juniper (Juniperus ashei Small 1901), showing that anatomical constraints on axial conductance can lead to significant trade-offs in woody roots as well. Root system water uptake responds to axial transport and can be limited by it in a biologically meaningful way.

Elevational trends in hydraulic efficiency and safety of Pinus cembra roots.

In alpine regions, elevational gradients in environmental parameters are reflected by structural and functional changes in plant traits. Elevational changes in plant water relations have also been demonstrated, but comparable information on root hydraulics is generally lacking. We analyzed the hydraulic efficiency (specific hydraulic conductivity ks, entire root system conductance KR) and vulnerability to drought-induced embolism (water potential at 50 % loss of conductivity Ψ50) of the roots of Pinus cembra trees growing along an elevational transect of 600 m. Hydraulic parameters of the roots were compared with those of the stem and related to anatomical traits {mean conduit diameter (d), wall reinforcement [(t/b)2]}. We hypothesized that temperature-related restrictions in root function would cause a progressive limitation of hydraulic efficiency and safety with increasing elevation. We found that both root ks and KR decreased from low (1600 m a.s.l.: ks 5.6 ± 0.7 kg m−1 s−1 MPa−1, KR 0.049 ± 0.005 kg m−2 s −1 MPa−1) to high elevation (2100 m a.s.l.: ks 4.2 ± 0.6 kg m−1 s−1 MPa−1, KR 0.035 ± 0.006 kg m−2 s−1 MPa−1), with small trees showing higher KR than large trees. ks was higher in roots than in stems (0.5 ± 0.05 kg m−1s−1MPa−1). Ψ50 values were similar across elevations and overall less negative in roots (Ψ50 −3.6 ± 0.1 MPa) than in stems (Ψ50 −3.9 ± 0.1 MPa). In roots, large-diameter tracheids were lacking at high elevation and (t/b)2 increased, while d did not change. The elevational decrease in root hydraulic efficiency reflects a limitation in timberline tree hydraulics. In contrast, hydraulic safety was similar across elevations, indicating that avoidance of hydraulic failure is important for timberline trees. As hydraulic patterns can only partly be explained by the anatomical parameters studied, limitations and/or adaptations at the pit level are likely.

Seasonal changes of whole root system conductance by a drought-tolerant grape root system

The role of root systems in drought tolerance is a subject of very limited information compared with above-ground responses. Adjustments to the ability of roots to supply water relative to shoot transpiration demand is proposed as a major means for woody perennial plants to tolerate drought, and is often expressed as changes in the ratios of leaf to root area (AL:AR). Seasonal root proliferation in a directed manner could increase the water supply function of roots independent of total root area (AR) and represents a mechanism whereby water supply to demand could be increased. To address this issue, seasonal root proliferation, stomatal conductance (gs) and whole root system hydraulic conductance (kr) were investigated for a drought-tolerant grape root system (Vitis berlandieri×V. rupestris cv. 1103P) and a non-drought-tolerant root system (Vitis riparia×V. rupestris cv. 101-14Mgt), upon which had been grafted the same drought-sensitive clone of Vitis vinifera cv. Merlot. Leaf water potentials (ψL) for Merlot grafted onto the 1103P root system (–0.91±0.02 MPa) were +0.15 MPa higher than Merlot on 101-14Mgt (–1.06±0.03 MPa) during spring, but dropped by approximately –0.4 MPa from spring to autumn, and were significantly lower by –0.15 MPa (–1.43±0.02 MPa) than for Merlot on 101-14Mgt (at –1.28±0.02 MPa). Surprisingly, gs of Merlot on the drought-tolerant root system (1103P) was less down-regulated and canopies maintained evaporative fluxes ranging from 35–20 mmol vine−1 s−1 during the diurnal peak from spring to autumn, respectively, three times greater than those measured for Merlot on the drought-sensitive rootstock 101-14Mgt. The drought-tolerant root system grew more roots at depth during the warm summer dry period, and the whole root system conductance (kr) increased from 0.004 to 0.009 kg MPa−1 s−1 during that same time period. The changes in kr could not be explained by xylem anatomy or conductivity changes of individual root segments. Thus, the manner in which drought tolerance was conveyed to the drought-sensitive clone appeared to arise from deep root proliferation during the hottest and driest part of the season, rather than through changes in xylem structure, xylem density or stomatal regulation. This information can be useful to growers on a site-specific basis in selecting rootstocks for grape clonal material (scions) grafted to them.

Rapid flood-induced stomatal closure accompanies xylem sap transportation of root-derived acetaldehyde and ethanol in Forsythia

Plants grown in containers frequently suffer from difficulties in managing their water status due to either insufficient or too much water. In the case of the latter, little information is available regarding how container-grown woody plants respond to anaerobic media. The aims of this work were therefore to use Forsythia as a model woody plant system to provide a mechanistic understanding of the physiological events and their timing during soil flooding. Exposure of pot-grown Forsythia to root hypoxia had a dramatic effect on leaf growth and stomatal conductance. Within 24 h of flooding a decline in leaf growth rate was detected along with a reduction in stomatal conductance. The effects of hypoxia appear initially with older leaves, but if flooding is prolonged (>2 days) younger expanding leaves are affected. These responses and their timing have not been described for woody perennial plants but appear comparable to those described for herbaceous plants such as tomato and castor bean. Measurements of stem and leaf tissue and during flooding showed large time dependent increases in the concentrations of acetaldehyde and ethanol; products associated with anaerobic respiration. Both of these chemicals were shown to be root-derived and are produced in a significant amount only as flooding time increases and the decline in leaf growth and conductance become apparent. Xylem sap was collected at a range of flow rates to measure whole root system conductance and determine changes in delivery of sap flux constituents. Calculated delivery rates of acetaldehyde and ethanol changed little with sap flow, particularly in well-drained control plants, while root hydraulic conductance declined when measured, 4 days after flooding. However, neither acetaldehyde nor ethanol, when used in a detached leaf transpiration bioassay, at physiologically realist concentrations (as determined from sap collection) failed to induce a dramatic reduction in leaf transpiration rates. The reasons for this discrepancy are discussed.

Wheat Plant Hydraulic Properties Under Prolonged Experimental Drought: Stronger Decline in Root-system Conductance than in Leaf Area

Wheat plants (Triticum aestivum var. INTA x2018;Cinco Cerros’) were grown in pots with fine sand under a rain-out shelter to assess their response to a water shortage spanning most of the growth cycle. Three watering treatments, based on different thresholds of plant-available water, were started 8 weeks after sowing and maintained for 10 weeks. After allowing recovery from any short-term embolism, stem-segment and root-system hydraulic conductances were then measured by standard low-pressure methods. Stress treatments reduced, as compared to controls, tiller number (by 31% and 41% for moderate and intense drought, respectively), total plant biomass (by 21% and 52%) and total plant leaf area (43% and 68%). The capacity of stems to transport water was reduced only by the most intense treatment (and then by no more than 50%), but root-system hydraulic conductance (k R) was strongly reduced by both treatments (37% and 80%, respectively). The transport capacity of belowground structures decreased not only on an absolute basis (k R), but also per unit root mass (K RS: 51% and 83%) and per unit of leaf area (K RL: 23% and 73%). Simulation of maximum transpiration under different soil and plant water conditions indicate that these changes in plant hydraulics had a significant impact on either transpiration at the leaf level or leaf water status for a given transpiration rate.

Changes in Root Hydraulic Conductivity During Wheat Evolution

Abstract: A better understanding of the mechanisms of water uptake by plant roots should be vital for improving drought resistance and water use efficiency (WUE). In the present study, we have demonstrated correlations between root system hydraulic conductivity and root characteristics during evolution using six wheat evolution genotypes (solution culture) with different ploidy chromosome sets (Triticum boeoticum Bioss., T. monococcum L.: 2n=2x=14; T. dicoccides Koern., T. dicoccon (Schrank) Schuebl.: 2n = 4x = 28; T. vulgare Vill., T. aestivum L. cv. Xiaoyan No. 6: 2n = 6x = 42). The experimental results showed that significant correlations were found between root system hydraulic conductivity and root characteristics of the materials with the increase in ploidy chromosomes (2x→6x) during wheat evolution. Hydraulic conductivity of the wheat root system at the whole‐plant level was increased with chromosome ploidy during evolution, which was positively correlated with hydraulic conductivity of single roots, whole plant biomass, root average diameter, and root growth (length, area), whereas the root/shoot ratio had an inverse correlation with the hydraulic conductivity of root system with increasing chromosome ploidy during wheat evolution. Therefore, it is concluded that that the water uptake ability of wheat roots was strengthened from wild to modern cultivated species during evolution, which will provide scientific evidence for genetic breeding to improve the WUE of wheat by genetic engineering.(Managing editor: Ya‐Qin HAN)

Relationships between Root System Water Transport Properties and Plant Size in Phaseolus

Root system hydraulic conductivity (L(P)) was measured on Phaseolus plants of different ages and sizes. Data analysis showed that L(P) changed in a complex manner depending on plant size. As the plants increased in size, L(P) increased initially then gradually decreased followed by a final modest increase. Values for L(P) ranged between 0.8 x 10(-6) and 6.1 x 10(-6) centimeter per second per bar. Relationships between the root flow per unit leaf area at a pressure differential of 3 bars (QPL(3)), as well as the total root system conductance (L(R)), and plant size were also examined. Values for QPL(3) varied with plant size, somewhat like L(P). L(R) values continuously increased with plant size at rates which depended on the growth rate of the root surface area as well as L(P). Comparison of our data with the root conductivity constant (k(r)) of Taylor and Klepper (1975 Soil Sci, 120: 57-67) showed good agreement. The observations on Phaseolus were also confirmed for Glycine. Values for L(P) and k(r) of both species were within the same range.