Simulation of root water uptake under consideration of nonequilibrium dynamics in the rhizosphere

The narrow region of soil around roots, the so-called rhizosphere, defers in its hydraulic properties from the bulk soil. The rhizosphere hydraulic properties primarily depend on the drying and wetting rate of mucilage, a polymeric gel exuded by plant roots. Under equilibrium conditions mucilage increases the water holding capacity. Upon drying mucilage turns hydrophobic and makes the rhizosphere temporarily water repellent. There are several models of root water uptake, from analytical models of water flow to a single root to complex numerical models that consider the root architecture. Most of these models, however, do not account for the specific hydraulic properties of the rhizosphere. Here we describe a single-root model that includes the altered hydraulic properties of the rhizosphere due to mucilage exudation. We use the model to reproduce existing experiments reporting unexpected and puzzling hysteresis in the rhizosphere, which could not be explained under the assumption of homogeneous hydraulic properties. In our model the hydraulic properties depend on the concentration of mucilage. This enables a continuous transition from the bulk soil to the root surface. We assumed that: (a) mucilage increases the water holding capacity in equilibrium conditions, (b) hydrophobicity, swelling and shrinking of mucilage cause a nonequilibrium relation between water content and water potential and (c) mucilage reduces the mobility of water molecules in the liquid phase resulting in a lower hydraulic conductivity at a given water content. Our model reproduces well the experiments and suggests that mucilage softens drought stress in plants during severe drying events.