The junction between root and soil is more than just two surfaces touching. It is often marked by a mucilaginous layer, rich in bacteria and fungi, that may be a few hundred micrometres thick, and in which adjacent soil particles are half or even wholly embedded. This is called the rhizosphere (Chapter 4). The hydraulic conductivity of this layer is not known, but there are two properties of this interfacial zone that may influence the flow of water across it. The first is that extensive gaps may occur within it, which strongly impede the flow. The second is that any exclusion of ions from the soil solution may result in large increases in the concentrations of these ions at the surface of the roots and this, because of the osmotic effects, would also impede the flow of
Water usually flows from soil to root as liquid, carrying with it dissolved nutrients. But there may be times when the hydraulic continuity between root and soil is so poor that a substantial proportion of the flow of water is as vapour. So far, nobody has managed to measure such flows, or even the relative importance of liquid and vapour flow, but their possible importance can be appreciated with the help of a few
The flow of vapour through soil is by diffusion, in contrast to that of liquid water, which moves in bulk. The consequence of this, and the fact that the vapour pressure of water is close to saturated in moist soil so that gradients in it are slight, is that water flows as vapour many times more slowly, in response to a given difference in water potential, than it does as
There are two main reasons why a root may be in poor hydraulic contact with the soil. The first is that it may be growing in a pre-existing pore larger than itself, so that it makes only glancing contact with the wall of the pore (see Figure 3.42 above). The second is that it has shrunk within a pore that it had itself made and which it had previously fitted snugly. What would induce a root to shrink? A fall in the water potential of the cortex would do so, for the cells there are thin-walled and separated in part by intercellular gas spaces, so they are likely to buckle. Observations made in rhizotrons (glass-walled tunnels for observing the behaviour of roots in the field) clearly show a diurnal shrinkage in cotton roots of up to 40% where the roots are growing in large pores, but we still do not know, because they are so difficult to observe, whether roots growing in intimate contact with the soil are similarly prone to shrink. A few observations made using neutron autoradiography have shown no shrinkage in roots growing in the
Because of the small size of the interfacial zone, it is very difficult to measure concentrations of ions within it. Rough estimates have been made by carefully separating the roots from the bulk of the soil, leaving only tenacious soil attached, then shaking off this remaining soil and analysing it. Such analyses do show somewhat higher concentrations (compared with those in the bulk soil) of ions such as sodium that we would expect to be largely excluded by the roots. But because the gradients in concentration are likely to be very large close to the root, these analyses are hard to assess for their consequences on the water relations of the plant. The gradients are likely to be especially large when the soil is becoming fairly dry, for the diffusion coefficient for the solutes falls by a few orders of magnitude as the soil dries, so that any excluded solutes will diffuse away from the root surface only very slowly.