Structures in the cell wall that enable hygroscopic movement of wheat awns

Abstract

The dispersal unit of wild wheat bears two prominent filaments called awns. The awns bend as they dry and straighten in a damp environment. This hygroscopic movement is explained by the orientation of the cellulose fibrils that build the cell wall, as follows. The stiff fibrils are embedded in a soft hygroscopic matrix. When the cell wall dries, the matrix shrinks but the fibrils do not. Therefore, the cell wall contracts in a direction perpendicular to the fibril orientation. Using X-ray scattering we identified a region at the base of the awn that contains fibrils aligned in all directions. This is the active part, which contracts as it dries and pulls the awn to a bent position. Cryo-scanning electron microscopy revealed sequential laminas which are rotated to form a nano-scale plywood construction, implying planar local order within the global isotropy. Water molecules absorbed into the matrix probably cause large microscopic distortions by expanding neighboring layers in perpendicular directions. This is thought to cause opening of tiny gaps between fiber layers, to facilitate the exchange and the transport of water through the cell wall, and thereby to increase the sensitivity of the actuating unit to moderate changes in humidity.