The rhizosphere is the zone of soil influenced by a plant root and is critical for plant health and nutrient acquisition. All below ground resources must pass through this dynamic zone prior to their capture by plant roots. However, researching the undisturbed rhizosphere has proved very challenging. Here we compare the temporal changes to the intact rhizosphere pore structure during the emergence of a developing root system in different soils. High resolution X-ray Computed Tomography (CT) was used to quantify the impact of root development on soil structural change, at scales relevant to individual micro-pores and aggregates (µm). A comparison of micro-scale structural evolution in homogenously packed soils highlighted the impacts of a penetrating root system in changing the surrounding porous architecture and morphology. Results indicate the structural zone of influence of a root can be more localised than previously reported (µm scale rather than mm scale). With time, growing roots significantly alter the soil physical environment in their immediate vicinity through reducing root-soil contact and crucially increasing porosity at the root-soil interface and not the converse as has often been postulated. This ‘rhizosphere pore structure’ and its impact on associated dynamics are discussed.

In this review, we examine how roots penetrate a structured soil. We first examine the relationship between soil water status and its mechanical strength, as well as the ability of the soil to supply water to the root. We identify these as critical soil factors, because it is primarily in drying soil that mechanical constraints limit root elongation. Water supply to the root is important because root water status affects growth pressures and root stiffness. To simplify the bewildering complexity of soil-root interactions, the discussion is focused around the special cases of root elongation in soil with pores much smaller than the root diameter and the penetration of roots at interfaces within the soil. While it is often assumed that the former case is well understood, many unanswered questions remain. While low soil-root friction is often viewed as a trait conferring better penetration of strong soils, it may also increase the axial pressure on the root tip and in so doing reduce the rate of cell division and/or expansion. The precise trade-off between various root traits involved in root elongation in homogeneous soil remains to be determined. There is consensus that the most important factors determining root penetration at an interface are the angle at which the root attempts to penetrate the soil, root stiffness, and the strength of the soil to be penetrated. The effect of growth angle on root penetration implicates gravitropic responses in improved root penetration ability. Although there is no work that has explored the effect of the strength of the gravitropic responses on penetration of hard layers, we attempt to outline possible interactions. Impacts of soil drying and strength on phytohormone concentrations in roots, and consequent root-to-shoot signalling, are also considered.

Background and aims There is an urgent need to develop new high throughput approaches to phenotype roots in the field. Excavating roots to make direct measurements is labour intensive. An alternative to excavation is to measure soil drying profiles and to infer root activity. Methods We grew 23 lines of wheat in 2013, 2014 and 2015. In each year we estimated soil water profiles with electrical resistance tomography (ERT), electromagnetic inductance (EMI), penetrometer measurements and measurements of soil water content. We determined the relationships between the measured variable and soil water content and matric potential. Results We found that ERT and penetrometer measurements were closely related to soil matric potential and produced the best discrimination between wheat lines.We found genotypic differences in depth of water uptake in soil water profiles and in the extent of surface drying. Conclusions Penetrometer measurements can provide a reliable approach to comparing soil drying profiles by different wheat lines, and genotypic rankings are repeatable across years. EMI, which is more sensitive to soil water content than matric potential, and is less effective in drier soils than the penetrometer or ERT, nevertheless can be used to rapidly screen large populations for differences in root activity.

In this opinion article we examine the relationship between penetrometer resistance and soil depth in the field. Assuming that root growth is inhibited at penetrometer resistances > 2.5 MPa, we conclude that in most circumstances the increases in penetrometer resistance with depth are sufficiently great to confine most deep roots to elongating in existing structural pores. We suggest that deep rooting is more likely related to the interaction between root architecture and soil structure than it is to the ability of a root to deform strong soil. Although the ability of roots to deform strong soil is an important trait, we propose it is more closely related to root exploration of surface layers than deep rooting.

HighlightsWe present evidence to support the hypothesis that the general and well-documented shape of the relationship between root length density and soil depth in UK grown winter wheat is related to the increase in soil strength with depth.Effects of the soil environment on root length distribution were greater than genetic effects and this was most likely related to soil saturation.In a dry season, there was genotypic variation in rooting depth.Greater root length at depth in the dwarf NILs suggests that deep rooting is not simply related to plant height.