Impact of root hairs on microscale soil physical properties in the field.

Aims: Recent laboratory studies revealed that root hairs may alter soil physical behaviour, influencing soil porosity and water retention on the small scale. However, the results are not consistent, and it is not known if structural changes at the small-scale have impacts at larger scales. Therefore, we evaluated the potential effects of root hairs on soil hydro-mechanical properties in the field using rhizosphere-scale physical measurements. Methods: Changes in soil water retention properties as well as mechanical and hydraulic characteristics were monitored in both silt loam and sandy loam soils. Measurements were taken from plant establishment to harvesting in field trials, comparing three barley genotypes representing distinct phenotypic categories in relation to root hair length. Soil hardness and elasticity were measured using a 3-mm-diameter spherical indenter, while water sorptivity and repellency were measured using a miniaturized infiltrometer with a 0.4-mm tip radius. Results: Over the growing season, plants induced changes in the soil water retention properties, with the plant available water increasing by 21%. Both soil hardness (P = 0.031) and elasticity (P = 0.048) decreased significantly in the presence of root hairs in silt loam soil, by 50% and 36%, respectively. Root hairs also led to significantly smaller water repellency (P = 0.007) in sandy loam soil vegetated with the hairy genotype (-49%) compared to the hairless mutant. Conclusions: Breeding of cash crops for improved soil conditions could be achieved by selecting root phenotypes that ameliorate soil physical properties and therefore contribute to increased soil health. Supplementary Information: The online version contains supplementary material available at 10.1007/s11104-022-05530-1.

Significance of root hairs for plant performance under contrasting field conditions and water deficit.

BACKGROUND AND AIMS: Previous laboratory studies have suggested selection for root hair traits in future crop breeding to improve resource use efficiency and stress tolerance. However, data on the interplay between root hairs and open-field systems, under contrasting soils and climate conditions, are limited. As such, this study aims to experimentally elucidate some of the impacts that root hairs have on plant performance on a field scale. METHODS: A field experiment was set up in Scotland for two consecutive years, under contrasting climate conditions and different soil textures (i.e. clay loam vs. sandy loam). Five barley (Hordeum vulgare) genotypes exhibiting variation in root hair length and density were used in the study. Root hair length, density and rhizosheath weight were measured at several growth stages, as well as shoot biomass, plant water status, shoot phosphorus (P) accumulation and grain yield. KEY RESULTS: Measurements of root hair density, length and its correlation with rhizosheath weight highlighted trait robustness in the field under variable environmental conditions, although significant variations were found between soil textures as the growing season progressed. Root hairs did not confer a notable advantage to barley under optimal conditions, but under soil water deficit root hairs enhanced plant water status and stress tolerance resulting in a less negative leaf water potential and lower leaf abscisic acid concentration, while promoting shoot P accumulation. Furthermore, the presence of root hairs did not decrease yield under optimal conditions, while root hairs enhanced yield stability under drought. CONCLUSIONS: Selecting for beneficial root hair traits can enhance yield stability without diminishing yield potential, overcoming the breeder's dilemma of trying to simultaneously enhance both productivity and resilience. Therefore, the maintenance or enhancement of root hairs can represent a key trait for breeding the next generation of crops for improved drought tolerance in relation to climate change.

Mechanisms of root reinforcement in soils: an experimental methodology using four-dimensional X-ray computed tomography and digital volume correlation.

Vegetation on railway or highway slopes can improve slope stability through the generation of soil pore water suctions by plant transpiration and mechanical soil reinforcement by the roots. To incorporate the enhanced shearing resistance and stiffness of root-reinforced soils in stability calculations, it is necessary to understand and quantify its effectiveness. This requires integrated and sophisticated experimental and multi-scale modelling approaches to develop an understanding of the processes at different length scales, from individual root-soil interaction through to full soil-profile or slope scale. One of the challenges with multi-scale models is ensuring that they sufficiently closely represent real behaviour. This requires calibration against detailed high-quality and data-rich experiments. This study presents a novel experimental methodology, which combines in situ direct shear loading of a willow root-reinforced soil with X-ray computed tomography to capture the three-dimensional chronology of soil and root deformation within the shear zone. Digital volume correlation (DVC) analysis was applied to the computed tomography dataset to obtain full-field three-dimensional displacement and strain information. This paper demonstrates the feasibility and discusses the challenges associated with DVC experiments on root-reinforced soils.

Significance of root hairs at the field scale - modelling root water and phosphorus uptake under different field conditions.

BACKGROUND AND AIMS: Root hairs play a significant role in phosphorus (P) extraction at the pore scale. However, their importance at the field scale remains poorly understood. METHODS: This study uses a continuum model to explore the impact of root hairs on the large-scale uptake of P, comparing root hair influence under different agricultural scenarios. High vs low and constant vs decaying P concentrations down the soil profile are considered, along with early vs late precipitation scenarios. RESULTS: Simulation results suggest root hairs accounted for 50% of total P uptake by plants. Furthermore, a delayed initiation time of precipitation potentially limits the P uptake rate by over 50% depending on the growth period. Despite the large differences in the uptake rate, changes in the soil P concentration in the domain due to root solute uptake remains marginal when considering a single growth season. However, over the duration of 6 years, simulation results showed that noticeable differences arise over time. CONCLUSION: Root hairs are critical to P capture, with uptake efficiency potentially enhanced by coordinating irrigation with P application during earlier growth stages of crops.

Surface tension, rheology and hydrophobicity of rhizodeposits and seed mucilage influence soil water retention and hysteresis.

AIMS: Rhizodeposits collected from hydroponic solutions with roots of maize and barley, and seed mucilage washed from chia, were added to soil to measure their impact on water retention and hysteresis in a sandy loam soil at a range of concentrations. We test the hypothesis that the effect of plant exudates and mucilages on hydraulic properties of soils depends on their physicochemical characteristics and origin. METHODS: Surface tension and viscosity of the exudate solutions were measured using the Du Noüy ring method and a cone-plate rheometer, respectively. The contact angle of water on exudate treated soil was measured with the sessile drop method. Water retention and hysteresis were measured by equilibrating soil samples, treated with exudates and mucilages at 0.46 and 4.6 mg g-1 concentration, on dialysis tubing filled with polyethylene glycol (PEG) solution of known osmotic potential. RESULTS: Surface tension decreased and viscosity increased with increasing concentration of the exudates and mucilage in solutions. Change in surface tension and viscosity was greatest for chia seed exudate and least for barley root exudate. Contact angle increased with increasing maize root and chia seed exudate concentration in soil, but not barley root. Chia seed mucilage and maize root rhizodeposits enhanced soil water retention and increased hysteresis index, whereas barley root rhizodeposits decreased soil water retention and the hysteresis effect. The impact of exudates and mucilages on soil water retention almost ceased when approaching wilting point at -1500 kPa matric potential. CONCLUSIONS: Barley rhizodeposits behaved as surfactants, drying the rhizosphere at smaller suctions. Chia seed mucilage and maize root rhizodeposits behaved as hydrogels that hold more water in the rhizosphere, but with slower rewetting and greater hysteresis.

Morphological and genetic characterisation of the root system architecture of selected barley recombinant chromosome substitution lines using an integrated phenotyping approach.

Discoveries on the genetics of resource acquisition efficiency are limited by the ability to measure plant roots in sufficient number and with adequate genotypic variability. This paper presents a root phenotyping study that explores ways to combine live imaging and computer algorithms for model-based extraction of root growth parameters. The study is based on a subset of barley Recombinant Chromosome Substitution Lines (RCSLs) and a combinatorial approach was designed for fast identification of the regions of the genome that contribute the most to variations in root system architecture (RSA). Results showed there was a strong genotypic variation in root growth parameters within the set of genotypes studied. The chromosomal regions associated with primary root growth differed from the regions of the genome associated with changes in lateral root growth. The concepts presented here are discussed in the context of identifying root QTL and its potential to assist breeding for novel crops with improved root systems.

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 6 µm 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.

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

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. HIGHLIGHTS: Rheological 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.

Can root electrical capacitance be used to predict root mass in soil?

BACKGROUND: Electrical capacitance, measured between an electrode inserted at the base of a plant and an electrode in the rooting substrate, is often linearly correlated with root mass. Electrical capacitance has often been used as an assay for root mass, and is conventionally interpreted using an electrical model in which roots behave as cylindrical capacitors wired in parallel. Recent experiments in hydroponics show that this interpretation is incorrect and a new model has been proposed. Here, the new model is tested in solid substrates. METHODS: The capacitances of compost and soil were determined as a function of water content, and the capacitances of cereal plants growing in sand or potting compost in the glasshouse, or in the field, were measured under contrasting irrigation regimes. KEY RESULTS: Capacitances of compost and soil increased with increasing water content. At water contents approaching field capacity, compost and soil had capacitances at least an order of magnitude greater than those of plant tissues. For plants growing in solid substrates, wetting the substrate locally around the stem base was both necessary and sufficient to record maximum capacitance, which was correlated with stem cross-sectional area: capacitance of excised stem tissue equalled that of the plant in wet soil. Capacitance measured between two electrodes could be modelled as an electrical circuit in which component capacitors (plant tissue or rooting substrate) are wired in series. CONCLUSIONS: The results were consistent with the new physical interpretation of plant capacitance. Substrate capacitance and plant capacitance combine according to standard physical laws. For plants growing in wet substrate, the capacitance measured is largely determined by the tissue between the surface of the substrate and the electrode attached to the plant. Whilst the measured capacitance can, in some circumstances, be correlated with root mass, it is not a direct assay of root mass.

Root border cells take up and release glucose-C.

BACKGROUND AND AIMS: Border cells are released from the root tips of many plant species, and can remain viable in the rhizosphere for 1 week. Whether border cells are capable of controlled glucose exchange with their environment was investigated. METHODS: Border cells were removed from Zea mays L. root tips, and immersed in (14)C-labelled D-glucose. In one experiment, the hexose transport inhibitor, phlorizin, was used to investigate active glucose uptake from a range of glucose concentrations. In another experiment, glucose efflux from border cells was monitored over time. KEY RESULTS: Glucose uptake by the border cells increased with increasing glucose concentration from 0.2 to 20 mm. At 0.2 mm glucose, uptake was mainly active, as evidenced by the approx. 60 % inhibition with phlorizin. At 2 and 20 mm glucose, however, uptake was mainly via diffusion, as phlorizin inhibition was negligible. Glucose efflux increased with time for live border cells in both 2 and 20 mm glucose. There was no clear efflux/time pattern for heat-killed border cells. CONCLUSIONS: Border cells actively take up glucose, and also release it. Under our experimental conditions, glucose uptake and efflux were of similar order of magnitude. In the rhizosphere net glucose exchange will almost certainly depend on local soil conditions.

Does the Presence of Detached Root Border Cells of Zea mays Alter the Activity of the Pathogenic Nematode Meloidogyne incognita?

ABSTRACT The root-knot nematode Meloidogyne incognita is a major pathogen of a range of important crops. Currently, control is typically achieved by the use of nematicides. However, recent work suggests that manipulating the ability of roots to slough off border cells, which then act as a decoy to the nematode, can significantly decrease damage to the roots. We investigated the attractiveness of border cells to M. incognita and the response of the nematode to border cells in close proximity. We found very limited attraction, in that nematodes did not preferentially alter direction to move toward the border cells, but a large and significant increase in nematode speed was observed once they were in the immediate vicinity of border cells. We discuss the results in the context of physical and biological mechanisms in relation to the control of pathogenic nematodes.

Plant influence on rhizosphere hydraulic properties: direct measurements using a miniaturized infiltrometer.

• An infiltrometer device, 0.4 mm in radius was designed specifically to measure the hydraulic characteristics of rhizosphere soil. Its testing and application to the rhizosphere of four plant species-barley (Hordeum vulgare), oil-seed rape (Brassica napus), potato (Solanum tuberosum) and grass (Lolium multiflorum) - was described. • In excavated blocks of field soil, there was a significant influence of plant species on sorptivity and water repellency in the rhizosphere. • Further controlled laboratory tests on young plants in moist, sieved soil showed reduced water sorptivity owing to increased repellency in the rhizosphere compared with bulk soil for barley but not oil-seed rape. • Root exudates may clog pores or become hydrophobic on soil particle surfaces. The slightly higher water repellency measured in rhizosphere soil would have minimal influence on plant water uptake. However, it may provide a buffer against desiccation at lower water contents and reduce structural degradation of rhizosphere soil by slaking.

Plant roots release phospholipid surfactants that modify the physical and chemical properties of soil.

• Plant root mucilages contain powerful surfactants that will alter the interaction of soil solids with water and ions, and the rates of microbial processes. • The lipid composition of maize, lupin and wheat root mucilages was analysed by thin layer chromatography and gas chromatography-mass spectrometry. A commercially available phosphatidylcholine (lecithin), chemically similar to the phospholipid surfactants identified in the mucilages, was then used to evaluate its effects on selected soil properties. • The lipids found in the mucilages were principally phosphatidylcholines, composed mainly of saturated fatty acids, in contrast to the lipids extracted from root tissues. In soil at low tension, lecithin reduced the water content at any particular tension by as much as 10 and 50% in soil and acid-washed sand, respectively. Lecithin decreased the amount of phosphate adsorption in soil and increased the phosphate concentration in solution by 10%. The surfactant also reduced net rates of ammonium consumption and nitrate production in soil. • These experiments provide the first evidence we are aware of that plant-released surfactants will significantly modify the biophysical environment of the rhizosphere.