Water transport in plants: Mechanism of apparent changes in resistance during absorption

Leaf water potentials were measured at various rates of water absorption in whole plants and detached leaves of well-watered Helianthus annuus L. The experiments were conducted in the steady state, where changes in leaf hydration did not affect the measurements but both the transpiration and growth components of absorption could be observed. Calculations of the total plant resistance to water transport showed that the resistance at low fluxes was about 30 times the resistance at high fluxes. Most of the change took place in the leaves, since similar changes could be demonstrated in detached leaves. The roots accounted for little of the change, since they varied in resistance by a factor of only 2.5 as flow varied.To ascertain whether the protoplasts of the leaves varied in resistance by an amount which could account for the change in resistance to water transport, measurements of rates of water movement in and out of the protoplasm were made when gradients in water potential between the protoplasts and the water source were varied. These showed that water movement did not occur at rates which could account for high rates of transpiration even when large differences in potential drove flow. The high temperature sensitivity of efflux confirmed that the leaf protoplasts limited flow in these experiments. When the edge of the leaf was excised and flow occurred primarily through the vascular system of the leaf, the resistance was much lower than in the protoplasts. It is therefore concluded that the leaf protoplasts represent a high resistance to water transport and that a considerable portion of the water involved in transpiration must bypass them.Calculations based on a model of water transport showed that the protoplast resistance was almost 30 times larger than the resistance of the path leading from the soil to the leaf protoplasts. The decrease in resistance of the leaves with increasing rates of absorption was therefore attributed to a decrease in water movement in and out of leaf cells, which involved a high resistance, and an increase in movement around the leaf protoplasts, which involved a low resistance. Since the experiments were conducted at the steady state, the high resistances were apparent at low rates of flow where only growth occurred, whereas the low resistances could be observed at high rates of flow because growth did not occur and flow consisted solely of transpiration. Because of the high resistance of the protoplast pathway, leaf water potentials were governed more by protoplast water movement than by transpiration over a considerable range of rates of water absorption. This may explain some of the differences in earlier work on leaf water potentials and water transport.