The mechanism of water absorption by roots I. Preliminary studies on the effects of hydrostatic pressure gradients

Abstract

During transpiration the hydrostatic tension which develops in the xylem conducting elements of the root draws water from the soil through the intervening tissues of the cortex, etc. It is uncertain whether this movement is entirely diffusional or in part a mass flow. To detect any such mass flow tomato plants grown in water culture were decapitated and placed in a canister through the lid of which the cut stem protruded and in which the pressure on the culture medium could be raised. The resulting rate of exudation (flux) was measured, and compared with the flux caused by an equivalent difference in osmotic potential obtained by measuring the ∆ f. p. of the medium and sap exuded. If these values of flux were equal, movement was by diffusion alone, but if pressure caused a greater flux, an additional mass flow was indicated. Preliminary experiments indicated a much greater flux in response to differences of pressure than osmotic potential, but accurate assessment of the effect was precluded by difficulties inherent in this straightforward approach. A less direct technique was therefore devised; the change in flux caused by changing the osmotic potential of the external medium (the hydrostatic pressure being maintained constant) was compared with the change in flux caused by changing the external pressure (the osmotic potential of the external medium being kept constant). The changes in flux were measured in such a way as to minimize changes in the osmotic potential in the xylem and in resistances to diffusion or mass flow respectively. In this way the change in flux per unit change in osmotic potential difference across the cortex (osmotic permeability coefficient,k0) and the change in flux per unit change in pressure difference across the cortex (pressure permeability coefficient,kp) could be compared under the same pressure gradient and in addition the effects of pressure gradients onk0could be studied. Thus, the effects of a pressure gradient on the diffusional movement of water could be assessed, as well as any mass flow component of the flux detected and measured.