Root hydraulic conductivity is a limiting factor along the water pathways between the soil and the leaf, and root radial conductivity is itself defined by cell-scale hydraulic properties and anatomical features. However, quantifying the influence of anatomical features on the radial conductivity remains challenging due to complex time-consuming experimental procedures. We present an open-source computational tool, the Generator of Root Anatomy in R (GRANAR; http://granar.github.io), that can be used to rapidly generate digital versions of contrasted monocotyledon root anatomical networks. GRANAR uses a limited set of root anatomical parameters, easily acquired with existing image analysis tools. The generated anatomical network can then be used in combination with hydraulic models to estimate the corresponding hydraulic properties. We used GRANAR to reanalyze large maize (Zea mays) anatomical datasets from the literature. Our model was successful at creating virtual anatomies for each experimental observation. We also used GRANAR to generate anatomies not observed experimentally over wider ranges of anatomical parameters. The generated anatomies were then used to estimate the corresponding radial conductivities with the hydraulic model MECHA (model of explicit cross-section hydraulic architecture). Our simulations highlight the large importance of the width of the stele and the cortex. GRANAR is a computational tool that generates root anatomical networks from experimental data. It enables the quantification of the effect of individual anatomical features on the root radial conductivity.
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.
Root hydraulic properties play a central role in the global water cycle, in agricultural systems productivity, and in ecosystem survival as they impact the canopy water supply. However, the existing experimental methods to quantify root hydraulic conductivities, such as the root pressure probing, are particularly challenging, and their applicability to thin roots and small root segments is limited. Therefore, there is a gap in methods enabling easy estimations of root hydraulic conductivities in diverse root types. Here, we present a new pipeline to quickly estimate root hydraulic conductivities across different root types, at high resolution along root axes. Shortly, free-hand root cross-sections were used to extract a selected number of key anatomical traits.We used these traits to parametrize the Generator of Root Anatomy in R (GRANAR) model to simulate root anatomical networks. Finally, we used these generated anatomical networks within the Model of Explicit Cross-section Hydraulic Anatomy (MECHA) to compute an estimation of the root axial and radial hydraulic conductivities (k x and k r , respectively). Using this combination of anatomical data and computational models, we were able to create a root hydraulic conductivity atlas at the root system level, for 14-day-old pot-grown Zea mays (maize) plants of the var. B73. The altas highlights the significant functional variations along and between different root types. For instance, predicted variations of radial conductivity along the root axis were strongly dependent on the maturation stage of hydrophobic barriers. The same was also true for the maturation rates of the metaxylem vessels. Differences in anatomical traits along and across root types generated substantial variations in radial and axial conductivities estimated with our novel approach. Our methodological pipeline combines anatomical data and computational models to turn root crosssection images into a detailed hydraulic atlas. It is an inexpensive, fast, and easily applicable investigation tool for root hydraulics that complements existing complex experimental methods. It opens the way to high-throughput studies on the functional importance of root types in plant hydraulics, especially if combined with novel phenotyping techniques such as laser ablation tomography.
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 nonlinear environment (soil, atmosphere), which makes unclear what impact of each single root system on root system functioning actually is. On the other end of the root system modelling continuum, macroscopic root system models associate to each root system instance 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 are compatible with macroscopic parameter measurements. The aim of this study was to bridge the gap between these two modelling approaches by providing a fast and reliable tool, which eventually can help performing plant virtual breeding.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, and a review of architectural and hydraulic parameters of maize.To illustrate the tool’s potential, we generated contrasted maize hydraulic architectures that we compared with architectural (root length density) and hydraulic (root system conductance) observations. Observed variability of these traits was well captured by model ensemble runs We also analyzed 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 parameters to vary for efficient virtual root system breeding. MARSHAL enables inverse optimisations, sensitivity analyses and virtual breeding of maize hydraulic root architecture. It is available as an R package, an RMarkdown pipeline, and a web application.One-sentence summaryWe developed a dynamic hydraulic-architectural model of the root system, parameterized for maize, to generate contrasted hydraulic architectures, compatible with field and lab observations and that can be further analyzed in soil-root system models for virtual breeding.Authors contributionsF.M., X.D., M.J. and G.L. designed the study and defined its scope; F.M. and G.L. developed the model while associated tools were created by A.H. and G.L.; F.M. ran the model simulations and analyzed the results together with M.J and G.L.; F.M. and M.J. wrote the first version of this manuscript; all co-authors critically revised it.
Root hydraulic conductivity is an important determinant of plant water uptake capacity. In particular, the root radial conductivity is often thought to be a limiting factor along the water pathways between the soil and the leaf. The root radial conductivity is itself defined by cell scale hydraulic properties and anatomical features. However, quantifying the influence of anatomical features on the radial conductivity remains challenging due to complex, and time-consuming, experimental procedures. We present a new computation tool, the Generator of Root ANAtomy in R (GRANAR) that can be used to rapidly generate digital versions of root anatomical networks. GRANAR uses a limited set of root anatomical parameters, easily acquired with existing image analysis tools. The generated anatomical network can then be used in combination with hydraulic models to estimate the corresponding hydraulic properties. Heymans et al. 2019 -GRANAR -A Generator of Root ANAtomy in R -Preprint We used GRANAR to re-analyse large maize (Zea mays) anatomical datasets from the literature. Our model was successful at creating virtual anatomies for each experimental observation. We also used GRANAR to generate anatomies not observed experimentally, over wider ranges of anatomical parameters. The generated anatomies were then used to estimate the corresponding radial conductivities with the hydraulic model MECHA. This enabled us to quantify the effect of individual anatomical features on the root radial conductivity. In particular, our simulations highlight the large importance of the width of the stele and the cortex. GRANAR is an open-source project available here: http://granar.github.io Keywords Root anatomy, radial conductivity, anatomical model, Zea mays , aerenchyma, reanalysis One-Sentence summary: Generator of Root ANAtomy in R (GRANAR) is a new open-source computational tool that can be used to rapidly generate digital versions of root anatomical networks.
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