How do roots elongate in a structured soil?

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.