Towards an active matter theory of plant morphogenesis

Plant morphogenesis relies on dynamic growth deformations at the cell and tissue scales driven by osmotic fluxes. A mechanistic understanding of this phenomenon demands a physical framework that integrates cell imbibition, tissue mechanics, and water fluxes, as well as their biophysical and molecular regulations, within a theory of plant active matter capturing the open-system and out-of-equilibrium properties of tissues. Building on historical insights into growth geometry, physics, and mechanics, combined with recent experimental results, we outline the key challenges in modelling plant growth and propose steps towards a unified physical theory of plant morphogenesis, in which biological regulation, mechanical forces, and water fluxes interact to shape biological form through the fundamental principles of living matter.