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.

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.

Brachiaria species influence nitrate transport in soil by modifying soil structure with their root system.

Leaching of nitrate from fertilisers diminishes nitrogen use efficiency (the portion of nitrogen used by a plant) and is a major source of agricultural pollution. To improve nitrogen capture, grasses such as brachiaria are increasingly used, especially in South America and Africa, as a cover crop, either via intercropping or in rotation. However, the complex interactions between soil structure, nitrogen and the root systems of maize and different species of forage grasses remain poorly understood. This study explored how soil structure modification by the roots of maize (Zea maize), palisade grass (Brachiaria brizantha cv. Marandu) and ruzigrass (Brachiaria ruziziensis) affected nitrate leaching and retention, measured via chemical breakthrough curves. All plants were found to increase the rate of nitrate transport suggesting root systems increase the tendency for preferential flow. The greater density of fine roots produced by palisade grass, subtly decreased nitrate leaching potential through increased complexity of the soil pore network assessed with X-ray Computed Tomography. A dominance of larger roots in ruzigrass and maize increased nitrate loss through enhanced solute flow bypassing the soil matrix. These results suggest palisade grass could be a more efficient nitrate catch crop than ruzigrass (the most extensively used currently in countries such as Brazil) due to retardation in solute flow associated with the fine root system and the complex pore network.

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.

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.

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.

Plant exudates improve the mechanical conditions for root penetration through compacted soils.

BACKGROUND AND AIM: Plant exudates greatly affect the physical behaviour of soil, but measurements of the impact of exudates on compression characteristics are missing. Our aim is to provide these data and explore how plant exudates may enhance the restructuring of compacted soils following cycles of wetting and drying. METHODS: Two soils were amended with Chia (Salvia hispanica) seed exudate at 5 concentrations, compacted in cores to 200 kPa stress (equivalent to tractor stress), equilibrated to -50 kPa matric potential, and then compacted to 600 kPa (equivalent to axial root stress) followed by 3 cycles of wetting and drying and recompression to 600 kPa at -50 kPa matric potential. Penetration resistance (PR), compression index (CC) and pore characteristics were measured at various steps. RESULTS: PR decreased and CC increased with increasing exudate concentration. At 600 kPa compression, 1.85 mg exudate g-1 soil increased CC from 0.37 to 0.43 for sandy loam soil and from 0.50 to 0.54 for clay loam soil. After 3 wetting-drying cycles the clay loam was more resillient than the sandy loam soil, with resilience increasing with greater exudate concentration. Root growth modelled on PR data suggested plant exudates significantly eased root elongation in soil. CONCLUSION: Plant exudates improve compression characteristics of soils, easing penetration and enhancing recovery of root induced soil compaction.

Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits.

Root elongation in drying soil is generally limited by a combination of mechanical impedance and water stress. Relationships between root elongation rate, water stress (matric potential), and mechanical impedance (penetration resistance) are reviewed, detailing the interactions between these closely related stresses. Root elongation is typically halved in repacked soils with penetrometer resistances >0.8-2 MPa, in the absence of water stress. Root elongation is halved by matric potentials drier than about -0.5 MPa in the absence of mechanical impedance. The likelihood of each stress limiting root elongation is discussed in relation to the soil strength characteristics of arable soils. A survey of 19 soils, with textures ranging from loamy sand to silty clay loam, found that ∼10% of penetration resistances were >2 MPa at a matric potential of -10 kPa, rising to nearly 50% >2 MPa at - 200 kPa. This suggests that mechanical impedance is often a major limitation to root elongation in these soils even under moderately wet conditions, and is important to consider in breeding programmes for drought-resistant crops. Root tip traits that may improve root penetration are considered with respect to overcoming the external (soil) and internal (cell wall) pressures resisting elongation. The potential role of root hairs in mechanically anchoring root tips is considered theoretically, and is judged particularly relevant to roots growing in biopores or from a loose seed bed into a compacted layer of soil.

Investigating the effects of anaerobic and aerobic post-treatment on quality and stability of organic fraction of municipal solid waste as soil amendment.

The use of OFMSW for biogas and compost production is considered as a sustainable strategy in saving valuable landfill space while producing valuable product for soil application. This study examines the effects of anaerobic and aerobic post-treatment of OFMSW on the stability of anaerobic digestate and compost and soil quality using seed germination tests. Anaerobic digestion of OFMSW was carried out for fifteen days after which the residual anaerobic digestate was subjected to aerobic post-treatment for seventy days. Seed germination tests showed that fresh feedstock and digestates collected during anaerobic digestion and during the early stages of aerobic post-treatment were phytotoxic. However, phytotoxic effects were not observed in soils amended with the fully stabilised anaerobic digestate compost, ADC. It was also found that seed germination increases with dilution and incubation time, suggesting that lower soil application rates and longer lag periods between soil application of ADC and planting can reduce the amount of biodegradable organics in the ADC, thus enhancing the benefits of ADC as soil amendment.

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.

Development of delta(15)N stratification of NO(3)(-) in soil profiles.

New evidence, obtained using a robust method for measuring the delta(15)N of NO(3)(-)-N in soil, is consistent with denitrification being the major determinant in the vertical distribution of NO(3)(-)-delta(15)N in soil profiles. These data also suggest that varying moisture regimes result in different effects of soil NO(3)(-)-N leaching on residual whole soil delta(15)N.

Changes to water repellence of soil caused by the growth of white-rot fungi: studies using a novel microcosm system.

A microcosm system is described which permits assessment of the progressive growth of filamentous fungi through soil. We report on its application to measure the effects of Coriolus versicolor and Phanerochaete chrysosporium upon the sorptivity and water repellence of a mineral soil, measured using a miniature infiltration device. Both fungal species caused moderate sub-critical repellence. Since the pore structure was unaffected, the repellence was probably due to hydrophobic substances of fungal origin. This is the first report of changes in soil repellence caused by the growth of potential xenobiotic bioremediating fungi. The potential consequences are discussed.