The effect of root hairs on root water uptake is determined by root-soil contact and root hair shrinkage.

The effect of root hairs on water uptake remains controversial. In particular, the key root hair and soil parameters that determine their importance have been elusive. We grew maize plants (Zea mays) in microcosms and scanned them using synchrotron-based X-ray computed microtomography. By means of image-based modelling, we investigated the parameters determining the effectiveness of root hairs in root water uptake. We explicitly accounted for rhizosphere features (e.g. root-soil contact and pore structure) and took root hair shrinkage of dehydrated root hairs into consideration. Our model suggests that > 85% of the variance in root water uptake is explained by the hair-induced increase in root-soil contact. In dry soil conditions, root hair shrinkage reduces the impact of hairs substantially. We conclude that the effectiveness of root hairs on root water uptake is determined by the hair-induced increase in root-soil contact and root hair shrinkage. Although the latter clearly reduces the effect of hairs on water uptake, our model still indicated facilitation of water uptake by root hairs at soil matric potentials from -1 to -0.1 MPa. Our findings provide new avenues towards a mechanistic understanding of the role of root hairs on water uptake.

Soil-plant hydraulics explain stomatal efficiency-safety tradeoff.

The efficiency-safety tradeoff has been thoroughly investigated in plants, especially concerning their capacity to transport water and avoid embolism. Stomatal regulation is a vital plant behaviour to respond to soil and atmospheric water limitation. Recently, a stomatal efficiency-safety tradeoff was reported where plants with higher maximum stomatal conductance (gmax ) exhibited greater sensitivity to stomatal closure during soil drying, that is, less negative leaf water potential at 50% gmax (ψgs50 ). However, the underlying mechanism of this gmax -ψgs50 tradeoff remains unknown. Here, we utilized a soil-plant hydraulic model, in which stomatal closure is triggered by nonlinearity in soil-plant hydraulics, to investigate such tradeoff. Our simulations show that increasing gmax is aligned with less negative ψgs50 . Plants with higher gmax (also higher transpiration) require larger quantities of water to be moved across the rhizosphere, which results in a precipitous decrease in water potential at the soil-root interface, and therefore in the leaves. We demonstrated that the gmax -ψgs50 tradeoff can be predicted based on soil-plant hydraulics, and is impacted by plant hydraulic properties, such as plant hydraulic conductance, active root length and embolism resistance. We conclude that plants may therefore adjust their growth and/or their hydraulic properties to adapt to contrasting habitats and climate conditions.

Stomatal closure during water deficit is controlled by below-ground hydraulics.

BACKGROUND AND AIMS: Stomatal closure allows plants to promptly respond to water shortage. Although the coordination between stomatal regulation, leaf and xylem hydraulics has been extensively investigated, the impact of below-ground hydraulics on stomatal regulation remains unknown. METHODS: We used a novel root pressure chamber to measure, during soil drying, the relation between transpiration rate (E) and leaf xylem water pressure (ψleaf-x) in tomato shoots grafted onto two contrasting rootstocks, a long and a short one. In parallel, we also measured the E(ψleaf-x) relation without pressurization. A soil-plant hydraulic model was used to reproduce the measurements. We hypothesize that (1) stomata close when the E(ψleaf-x) relation becomes non-linear and (2) non-linearity occurs at higher soil water contents and lower transpiration rates in short-rooted plants. KEY RESULTS: The E(ψleaf-x) relation was linear in wet conditions and became non-linear as the soil dried. Changing below-ground traits (i.e. root system) significantly affected the E(ψleaf-x) relation during soil drying. Plants with shorter root systems required larger gradients in soil water pressure to sustain the same transpiration rate and exhibited an earlier non-linearity and stomatal closure. CONCLUSIONS: We conclude that, during soil drying, stomatal regulation is controlled by below-ground hydraulics in a predictable way. The model suggests that the loss of hydraulic conductivity occurred in soil. These results prove that stomatal regulation is intimately tied to root and soil hydraulic conductances.

Stomatal closure of tomato under drought is driven by an increase in soil-root hydraulic resistance.

The fundamental question as to what triggers stomatal closure during soil drying remains contentious. Thus, we urgently need to improve our understanding of stomatal response to water deficits in soil and atmosphere. Here, we investigated the role of soil-plant hydraulic conductance (Ksp ) on transpiration (E) and stomatal regulation. We used a root pressure chamber to measure the relation between E, leaf xylem water potential (ψleaf-x ) and soil water potential (ψsoil ) in tomato. Additional measurements of ψleaf-x were performed with unpressurized plants. A soil-plant hydraulic model was used to simulate E(ψleaf-x ) for decreasing ψsoil . In wet soils, E(ψleaf-x ) had a constant slope, while in dry soils, the slope decreased, with ψleaf-x rapidly and nonlinearly decreasing for moderate increases in E. The ψleaf-x measured in pressurized and unpressurized plants matched well, which indicates that the shoot hydraulic conductance did not decrease during soil drying and that the decrease in Ksp is caused by a decrease in soil-root conductance. The decrease of E matched well the onset of hydraulic nonlinearity. Our findings demonstrate that stomatal closure prevents the drop in ψleaf-x caused by a decrease in Ksp and elucidate a strong correlation between stomatal regulation and belowground hydraulic limitation.

Investigating the root plasticity response of Centaurea jacea to soil water availability changes from isotopic analysis.

Root water uptake is a key ecohydrological process for which a physically based understanding has been developed in the past decades. However, due to methodological constraints, knowledge gaps remain about the plastic response of whole plant root systems to a rapidly changing environment. We designed a laboratory system for nondestructive monitoring of stable isotopic composition in plant transpiration of a herbaceous species (Centaurea jacea) and of soil water across depths, taking advantage of newly developed in situ methods. Daily root water uptake profiles were obtained using a statistical Bayesian multisource mixing model. Fast shifts in the isotopic composition of both soil and transpiration water could be observed with the setup and translated into dynamic and pronounced shifts of the root water uptake profile, even in well watered conditions. The incorporation of plant physiological and soil physical information into statistical modelling improved the model output. A simple exercise of water balance closure underlined the nonunique relationship between root water uptake profile on the one hand, and water content and root distribution profiles on the other, illustrating the continuous adaption of the plant water uptake as a function of its root hydraulic architecture and soil water availability during the experiment.

Going with the Flow: Multiscale Insights into the Composite Nature of Water Transport in Roots.

As water often limits crop production, a more complete understanding of plant water capture and transport is necessary. Here, we developed MECHA, a mathematical model that computes the flow of water across the root at the scale of walls, membranes, and plasmodesmata of individual cells, and used it to test hypotheses related to root water transport in maize (Zea mays). The model uses detailed root anatomical descriptions and a minimal set of experimental cell properties, including the conductivity of plasma membranes, cell walls, and plasmodesmata, which yield quantitative and scale-consistent estimations of water pathways and root radial hydraulic conductivity (k r). MECHA revealed that the mainstream hydraulic theories derived independently at the cell and root segment scales are compatible only if osmotic potentials within the apoplastic domains are uniform. The results suggested that the convection-diffusion of apoplastic solutes explained most of the offset between estimated k r in pressure clamp and osmotic experiments, while the contribution of water-filled intercellular spaces was limited. Furthermore, sensitivity analyses quantified the relative impact of cortex and endodermis cell conductivity of plasma membranes on root k r and suggested that only the latter contributed substantially to k r due to the composite nature of water flow across roots. The explicit root hydraulic anatomy framework brings insights into contradictory interpretations of experiments from the literature and suggests experiments to efficiently address questions pertaining to root water relations. Its scale consistency opens avenues for cross-scale communication in the world of root hydraulics.

Functional-structural root-system model validation using a soil MRI experiment.

For the first time, a functional-structural root-system model is validated by combining a tracer experiment monitored with magnetic resonance imaging and three-dimensional modeling of water and solute transport.

Connecting the dots between computational tools to analyse soil-root water relations.

In recent years, many computational tools, such as image analysis, data management, process-based simulation, and upscaling tools, have been developed to help quantify and understand water flow in the soil-root system, at multiple scales (tissue, organ, plant, and population). Several of these tools work together or at least are compatible. However, for the uninformed researcher, they might seem disconnected, forming an unclear and disorganized succession of tools. In this article, we show how different studies can be further developed by connecting them to analyse soil-root water relations in a comprehensive and structured network. This 'explicit network of soil-root computational tools' informs readers about existing tools and helps them understand how their data (past and future) might fit within the network. We also demonstrate the novel possibilities of scale-consistent parameterizations made possible by the network with a set of case studies from the literature. Finally, we discuss existing gaps in the network and how we can move forward to fill them.

Root type matters: measurement of water uptake by seminal, crown, and lateral roots in maize.

The ability of plants to take up water from the soil depends on both the root architecture and the distribution and evolution of the hydraulic conductivities among root types and along the root length. The mature maize (Zea mays L.) root system is composed of primary, seminal, and crown roots together with their respective laterals. Our understanding of root water uptake of maize is largely based on measurements of primary and seminal roots. Crown roots might have a different ability to extract water from the soil, but their hydraulic function remains unknown. The aim of this study was to measure the location of water uptake in mature maize and investigate differences between seminal, crown, and lateral roots. Neutron radiography and injections of deuterated water were used to visualize the root architecture and water transport in 5-week-old maize root systems. Water was mainly taken up by crown roots. Seminal roots and their laterals, which were the main location of water uptake in younger plants, made a minor contribution to water uptake. In contrast to younger seminal roots, crown roots were also able to take up water from their most distal segments. The greater uptake of crown roots compared with seminal roots is explained by their higher axial conductivity in the proximal parts and by the fact that they are connected to the shoot above the seminal roots, which favors the propagation of xylem tension along the crown roots. The deeper water uptake of crown roots is explained by their shorter and fewer laterals, which decreases the dissipation of water potential along the roots.

Increased contribution of wheat nocturnal transpiration to daily water use under drought.

Increasing evidence suggests that in crops, nocturnal water use could represent 30% of daytime water consumption, particularly in semi-arid and arid areas. This raises the questions of whether nocturnal transpiration rates (TRN ) are (1) less influenced by drought than daytime TR (TRD ), (2) increased by higher nocturnal vapor pressure deficit (VPDN ), which prevails in such environments and (3) involved in crop drought tolerance. In this investigation, we addressed those questions by subjecting two wheat genotypes differing in drought tolerance to progressive soil drying under two long-term VPDN regimes imposed under naturally fluctuating conditions. A first goal was to characterize the response curves of whole-plant TRN and TRN /TRD ratios to progressive soil drying. A second goal was to examine the effect of VPDN increase on TRN response to soil drying and on 13 other developmental traits. The study revealed that under drought, TRN was not responsive to progressive soil drying and - intriguingly - that TRN seemingly increased with drought under high VPDN consistently for the drought-sensitive genotype. Because TRD was concomitantly decreasing with progressive drought, this resulted in TRN representing up to 70% of TRD at the end of the drydown. In addition, under drought, VPDN increase was found not to influence traits such as leaf area or stomata density. Overall, those findings indicate that TRN contribution to daily water use under drought might be much higher than previously thought, that it is controlled by specific mechanisms and that decreasing TRN under drought might be a valuable trait for improving drought tolerance.

Root system markup language: toward a unified root architecture description language.

The number of image analysis tools supporting the extraction of architectural features of root systems has increased in recent years. These tools offer a handy set of complementary facilities, yet it is widely accepted that none of these software tools is able to extract in an efficient way the growing array of static and dynamic features for different types of images and species. We describe the Root System Markup Language (RSML), which has been designed to overcome two major challenges: (1) to enable portability of root architecture data between different software tools in an easy and interoperable manner, allowing seamless collaborative work; and (2) to provide a standard format upon which to base central repositories that will soon arise following the expanding worldwide root phenotyping effort. RSML follows the XML standard to store two- or three-dimensional image metadata, plant and root properties and geometries, continuous functions along individual root paths, and a suite of annotations at the image, plant, or root scale at one or several time points. Plant ontologies are used to describe botanical entities that are relevant at the scale of root system architecture. An XML schema describes the features and constraints of RSML, and open-source packages have been developed in several languages (R, Excel, Java, Python, and C#) to enable researchers to integrate RSML files into popular research workflow.

Estimation of the hydraulic conductivities of lupine roots by inverse modelling of high-resolution measurements of root water uptake.

Background and Aims Radial and axial hydraulic conductivities are key parameters for proper understanding and modelling of root water uptake. Despite their importance, there is limited experimental information on how the radial and axial hydraulic conductivities vary along roots growing in soil. Here, a new approach was introduced to estimate inversely the profile of hydraulic conductivities along the roots of transpiring plants growing in soil. Methods A three-dimensional model of root water uptake was used to reproduce the measured profile of root water uptake along roots of lupine plant grown in soil. The profile of fluxes was measured using a neutron radiography technique combined with injection of deuterated water as tracer. The aim was to estimate inversely the profiles of the radial and axial hydraulic conductivities along the roots. Key Results The profile of hydraulic conductivities along the taproot and the lateral roots of lupines was calculated using three flexible scenarios. For all scenarios, it was found that the radial hydraulic conductivity increases towards the root tips, while the axial conductivity decreases. Additionally, it was found that in soil with uniform water content: (1) lateral roots were the main location of root water uptake; (2) water uptake by laterals decreased towards the root tips due to the dissipation of water potential along the root; and (3) water uptake by the taproot was higher in the distal segments and was negligible in the proximal parts, which had a low radial conductivity. Conclusions The proposed approach allows the estimation of the root hydraulic properties of plants growing in soil. This information can be used in an advanced model of water uptake to predict the water uptake of different root types or different root architectures under varying soil conditions.

Plant water uptake in drying soils.

Over the last decade, investigations on root water uptake have evolved toward a deeper integration of the soil and roots compartment properties, with the goal of improving our understanding of water acquisition from drying soils. This evolution parallels the increasing attention of agronomists to suboptimal crop production environments. Recent results have led to the description of root system architectures that might contribute to deep-water extraction or to water-saving strategies. In addition, the manipulation of root hydraulic properties would provide further opportunities to improve water uptake. However, modeling studies highlight the role of soil hydraulics in the control of water uptake in drying soil and call for integrative soil-plant system approaches.

Fine-resolution profile-scale data to depict the impact of tillage treatment and machine traffic on agricultural soil structure and hydrologic properties.

This data article provides high spatial resolution (1 cm) datasets and related figures of the penetrometer resistance (PR) and soil bulk density (BD) data of nine agricultural 50 × 160 cm soil profiles exposed to three tillage treatments and including a wheel track. Soil treatments are moldboard plowing (MP), deep loosening (DL), and minimum tillage (MT). It also provides bulk density data, soil moisture content at various suctions and the parameters of van Genuchten's model for 27 soil cores, and saturated hydraulic conductivity (Ks) of 49 soil cores. Both sample sets were sampled to cover the profile heterogeneity in two agricultural plots subjected to moldboard plowing and minimum tillage. Examples of reuse potential include (i) the use of these spatially explicit data in studies seeking to understand better and integrate the effect of treatment and machine traffic-induced soil structure in soil hydraulic and soil physical quality, and (ii) the development of pedotransfer functions with data incorporating the soil structural heterogeneity. This Data in Brief article complements the companion paper by Alonso et al. (2021) "A hybrid method for characterizing tillage-induced soil physical quality at the profile scale with fine spatial detail" in Soil and Tillage Research[1].

How to Define the Appropriate Spatial Resolution of Root Segments When Solving Water Flow in Root System Hydraulic Architectures.

In this chapter, we discuss the issue of balance between spatial resolution and computational efficiency in the context of the R-SWMS model. Based on the equations governing the water fluxes within the model, we propose here an objective and quantitative criterion which can help fix root segment size to both minimize computational load and achieve simulation according to a given accuracy degree.

Investigating Soil-Root Interactions with the Numerical Model R-SWMS.

In this chapter, we present the Root and Soil Water Movement and Solute transport model R-SWMS, which can be used to simulate flow and transport in the soil-plant system. The equations describing water flow in soil-root systems are presented and numerical solutions are provided. An application of R-SWMS is then briefly discussed, in which we combine in vivo and in silico experiments in order to decrypt water flow in the soil-root domain. More precisely, light transmission imaging experiments were conducted to generate data that can serve as input for the R-SWMS model. These data include the root system architecture, the soil hydraulic properties and the environmental conditions (initial soil water content and boundary conditions, BC). Root hydraulic properties were not acquired experimentally, but set to theoretical values found in the literature. In order to validate the results obtained by the model, the simulated and experimental water content distributions were compared. The model was then used to estimate variables that were not experimentally accessible, such as the actual root water uptake distribution and xylem water potential.

Mechanistic modeling of pesticide uptake with a 3D plant architecture model.

Meaningful assessment of pesticide fate in soils and plants is based on fate models that represent all relevant processes. With mechanistic models, these processes can be simulated based on soil, substance, and plant properties. We present a mechanistic model that simulates pesticide uptake from soil and investigate how it is influenced, depending on the governing uptake process, by root and substance properties and by distributions of the substance and water in the soil profile. A new root solute uptake model based on a lumped version of the Trapp model (Trapp, 2000) was implemented in a coupled version of R-SWMS-ParTrace models for 3-D water flow and solute transport in soil and root systems. Solute uptake was modeled as two individual processes: advection with the transpiration stream and diffusion through the root membrane. We set up the model for a FOCUS scenario used in the European Union (EU) for pesticide registration. Considering a single vertical root and advective uptake only, the root hydraulic properties could be defined so that water and substance uptake and substance fate in soil showed a good agreement with the results of the 1D PEARL model, one of the reference models used in the EU for pesticide registration. Simulations with a complex root system and using root hydraulic parameters reported in the literature predicted larger water uptake from the upper root zone, leading to larger pesticide uptake when pesticides are concentrated in the upper root zone. Dilution of root water concentrations at the top root zone with water with low pesticide concentration taken up from the bottom of the root zone leads to larger uptake of solute when uptake was simulated as a diffusive process. This illustrates the importance of modeling uptake mechanistically and considering root and solute physical and chemical properties, especially when root-zone pesticide concentrations are non-uniform.