Chemical agents transported by xylem mass flow propagate variation potentials

Long-distance signalling is important for coordinating plant responses to the environment. Variation potentials (VPs) are a type of long-distance electrical signal that are generated in plants in response to wounding or flaming. Unlike self-propagating action potentials, VPs can be measured beyond regions of dead or chemically treated tissue that block signal generation, suggesting a different mode of propagation. Two alternative propagation mechanisms have been proposed: movement of a chemical agent and a pressure wave through the vasculature. Variants of these two signalling mechanisms have been suggested. Here, we use simple models of the underlying physical processes to evaluate and compare these predictions against independent data. Our models suggest that chemical diffusion and pressure waves are unlikely to capture existing data with parameters that are known from other sources. The previously discarded hypothesis of mass flow in the xylem transporting a chemical agent, however, is able to reproduce experimental propagation speeds for VPs. We therefore suggest that chemical agents transported by mass flow within the xylem are more likely than a pressure wave or chemical diffusion as a VP propagation mechanism. Understanding this mode of long-distance signalling within plants is important for unravelling how plants coordinate physiological responses via cell-to-cell communication.