Slow, fast and furious: understanding the physics of plant movements.

The ability of plants to move is central to many physiological processes from development to tropisms, from nutrition to reproduction. The movement of plants or plant parts occurs over a wide range of sizes and time scales. This review summarizes the main physical mechanisms plants use to achieve motility, highlighting recent work at the frontier of biology and physics on rapid movements. Emphasis is given to presenting in a single framework pioneering biological studies of water transport and growth with more recent physics research on poroelasticity and mechanical instabilities. First, the basic osmotic and hydration/dehydration motors are described that contribute to movement by growth and reversible swelling/shrinking of cells and tissues. The speeds of these water-driven movements are shown to be ultimately limited by the transport of water through the plant body. Some plant structures overcome this hydraulic limit to achieve much faster movement by using a mechanical instability. The principle is to impose an 'energy barrier' to the system, which can originate from geometrical constraint or matter cohesion, allowing elastic potential energy to be stored until the barrier is overcome, then rapidly transformed into kinetic energy. Three of these rapid motion mechanisms have been elucidated recently and are described here: the snapping traps of two carnivorous plants, the Venus flytrap and Utricularia, and the catapult of fern sporangia. Finally, movement mechanisms are reconsidered in the context of the timescale of important physiological processes at the cellular and molecular level.