Nicolai Koebernick

High-resolution synchrotron imaging shows that root hairs influence rhizosphere soil structure formation

Summary In this paper, we provide direct evidence of the importance of root hairs on pore structure development at the root–soil interface during the early stage of crop establishment. This was achieved by use of high‐resolution (c. 5 μm) synchrotron radiation computed tomography (SRCT) to visualise both the structure of root hairs and the soil pore structure in plant–soil microcosms. Two contrasting genotypes of barley (Hordeum vulgare), with and without root hairs, were grown for 8 d in microcosms packed with sandy loam soil at 1.2 g cm−3 dry bulk density. Root hairs were visualised within air‐filled pore spaces, but not in the fine‐textured soil regions. We found that the genotype with root hairs significantly altered the porosity and connectivity of the detectable pore space (> 5 μm) in the rhizosphere, as compared with the no‐hair mutants. Both genotypes showed decreasing pore space between 0.8 and 0.1 mm from the root surface. Interestingly the root‐hair‐bearing genotype had a significantly greater soil pore volume‐fraction at the root–soil interface. Effects of pore structure on diffusion and permeability were estimated to be functionally insignificant under saturated conditions when simulated using image‐based modelling.

Plant exudates may stabilize or weaken soil depending on species, origin and time

&NA; We hypothesized that plant exudates could either gel or disperse soil depending on their chemical characteristics. Barley (Hordeum vulgare L. cv. Optic) and maize (Zea mays L. cv. Freya) root exudates were collected using an aerated hydroponic method and compared with chia (Salvia hispanica L.) seed exudate, a commonly used root exudate analogue. Sandy loam soil was passed through a 500‐μm mesh and treated with each exudate at a concentration of 4.6 mg exudate g−1 dry soil. Two sets of soil samples were prepared. One set of treated soil samples was maintained at 4°C to suppress microbial processes. To characterize the effect of decomposition, the second set of samples was incubated at 16°C for 2 weeks at −30 kPa matric potential. Gas chromatography‐mass spectrometry (GC‐MS) analysis of the exudates showed that barley had the largest organic acid content and chia the largest content of sugars (polysaccharide‐derived or free), and maize was in between barley and chia. Yield stress of amended soil samples was measured by an oscillatory strain sweep test with a cone plate rheometer. When microbial decomposition was suppressed at 4°C, yield stress increased 20‐fold for chia seed exudate and twofold for maize root exudate compared with the control, whereas for barley root exudate decreased to half. The yield stress after 2 weeks of incubation compared with soil with suppressed microbial decomposition increased by 85% for barley root exudate, but for chia and maize it decreased by 87 and 54%, respectively. Barley root exudation might therefore disperse soil and this could facilitate nutrient release. The maize root and chia seed exudates gelled soil, which could create a more stable soil structure around roots or seeds. HighlightsRheological measurements quantified physical behaviour of plant exudates and effect on soil stabilization.Barley root exudates dispersed soil, which could release nutrients and carbon.Maize root and chia seed exudates had a stabilizing effect on soil.Physical engineering of soil in contact with plant roots depends on the nature and origin of exudates.

Fluid flow in porous media using image-based modelling to parametrize Richards' equation

The parameters in Richards equation are usually calculated from experimentally measured values of the soil–water characteristic curve and saturated hydraulic conductivity. The complex pore structures that often occur in porous media complicate such parametrization due to hysteresis between wetting and drying and the effects of tortuosity. Rather than estimate the parameters in Richards equation from these indirect measurements, image-based modelling is used to investigate the relationship between the pore structure and the parameters. A three-dimensional, X-ray computed tomography image stack of a soil sample with voxel resolution of 6u2009μm has been used to create a computational mesh. The Cahn–Hilliard–Stokes equations for two-fluid flow, in this case water and air, were applied to this mesh and solved using the finite-element method in COMSOL Multiphysics. The upscaled parameters in Richards equation are then obtained via homogenization. The effect on the soil–water retention curve due to three different contact angles, 0°, 20° and 60°, was also investigated. The results show that the pore structure affects the properties of the flow on the large scale, and different contact angles can change the parameters for Richards equation.

Measurement of micro-scale soil deformation around roots using four-dimensional synchrotron tomography and image correlation

This study applied time lapse (four-dimensional) synchrotron X-ray computed tomography to observe micro-scale interactions between plant roots and soil. Functionally contrasting maize root tips were repeatedly imaged during ingress into soil columns of varying water content and compaction. This yielded sequences of three-dimensional densiometric data, representing time-resolved geometric soil and root configurations at the micronmetre scale. These data were used as inputs for two full-field kinematic quantification methods, which enabled the analysis of three-dimensional soil deformation around elongating roots. Discrete object tracking was used to track rigid mineral grains, while continuum digital volume correlation was used to track grey-level patterns within local sub-volumes. These techniques both allowed full-field soil displacements to be quantified at an intra-rhizosphere spatial sampling scale of less than 300 µm. Significant differences in deformation mechanisms were identified around different phenotypes under different soil conditions. A uniquely strong contrast was observed between intact and de-capped roots grown in dry, compacted soil. This provides evidence that functional traits of the root cap significantly reduce the amount of soil disturbance per unit of root elongation, with this effect being particularly significant in drier soil.

Correlative visualization of root mucilage degradation using X-ray CT and NMRI

Root exudates are a crucial component of the rhizosphere. Often, they take a form of a gel exuded by the plant roots and are thought to influence the soil aggregation, root penetration into soil, soil nutrient availability, immobilization of toxic cations and microbial activity amongst other things. In addition, the capacity of exudates to store water makes the plants potentially less susceptive to drought. Major components of root exudates are high molecular weight organic compounds consisting of predominantly polysaccharides and proteins, which makes it challenging to visualize using current rhizosphere visualization techniques, such as X-ray computed tomography (CT). In this contribution, we use correlative X-ray CT (resolution ~20 uf06dm) in combination with Magnetic Resonance Imaging (MRI, resolution ~120 uf06dm) to set up groundwork to enable in situ visualization of mucilage in soil. This multimodal approach is necessary because mucilage density closely matches that of water. We use chia seeds as mucilage analogue, because it has been found to have a similar consistency to root mucilage. Moreover, to understand mucilage development in time, a series of samples made by chia seeds placed in different porous media were prepared. Structurally and chemically, mucilage breaks down towards a water-like substance over a course of two weeks. Depending on its relative concentration, these changes were found to be less dominant when seeds were mixed in porous media. Having set up the groundwork for correlative imaging of chia seeds in water and an artificial soil (Nafion and sand/beads) this enables us to expand this imaging to deal with plant root exudates under natural conditions.

The effect of root exudates on rhizosphere water dynamics

Most water and nutrients essential for plant growth travel across a thin zone of soil at the interface between roots and soil, termed the rhizosphere. Chemicals exuded by plant roots can alter the fluid properties, such as viscosity, of the water phase, potentially with impacts on plant productivity and stress tolerance. In this paper, we study the effects of plant exudates on the macroscale properties of water movement in soil. Our starting point is a microscale description of two fluid flow and exudate diffusion in a periodic geometry composed from a regular repetition of a unit cell. Using multiscale homogenization theory, we derive a coupled set of equations that describe the movement of air and water, and the diffusion of plant exudates on the macroscale. These equations are parametrized by a set of cell problems that capture the flow behaviour. The mathematical steps are validated by comparing the resulting homogenized equations to the original pore scale equations, and we show that the difference between the two models is ≲7% for eight cells. The resulting equations provide a computationally efficient method to study plant–soil interactions. This will increase our ability to predict how contrasting root exudation patterns may influence crop uptake of water and nutrients.

Root-soil contact dynamics of Vicia faba in sand

AimsRoot shrinkage in drying soil has been shown repeatedly. The aim of this study was to investigate the dynamics of root-soil contact and its relationship with plant water status during soil drying.MethodsThe development of root-soil contact of Vicia faba L. during a drying period was studied. Plants (Nu2009=u20094) were grown in cylinders filled with a sandy soil. Samples were repeatedly scanned with an X-ray CT scanner to visualize root-soil contact. Soil matric potential, transpiration rate, and stomatal conductance were measured daily.ResultsRoot-soil contact was lower in taproots than in lateral roots at any time. Transpiration rate and stomatal conductance decreased before roots started to shrink. Root-soil contact decreased significantly over the course of the drying period, starting at soil matric potentials below −20xa0kPa. Root shrinkage did not differ significantly between taproots and laterals.ConclusionsThis study confirms previous findings with Lupinus albus roots in that roots shrink after transpiration rate decreases. The dynamics of root shrinkage are governed by soil water availability and transpirational demand.

Imaging microstructure of the barley rhizosphere: particle packing and root hair influences

Soil adjacent to roots has distinct structural and physical properties from bulk soil, affecting water and solute acquisition by plants. Detailed knowledge on how root activity and traits such as root hairs affect the three-dimensional pore structure at a fine scale is scarce and often contradictory. Roots of hairless barley (Hordeum vulgare L. cv Optic) mutant (NRH) and its wildtype (WT) parent were grown in tubes of sieved (<250xa0μm) sandy loam soil under two different water regimes. The tubes were scanned by synchrotron-based X-ray computed tomography to visualise pore structure at the soil-root interface. Pore volume fraction and pore size distribution were analysed vs distance within 1xa0mm of the root surface. Less dense packing of particles at the root surface was hypothesised to cause the observed increased pore volume fraction immediately next to the epidermis. The pore size distribution was narrower due to a decreased fraction of larger pores. There were no statistically significant differences in pore structure between genotypes or moisture conditions. A model is proposed that describes the variation in porosity near roots taking into account soil compaction and the surface effect at the root surface.

Dataset for High-resolution synchrotron imaging shows that root hairs influence rhizosphere soil structure formation

Dataset supports:nKoebernick, N.et al (2017). High-resolution synchrotron imaging shows that root hairs influence rhizosphere soil structure formation. New Phytologist.