Three-dimensional printing, muscles, and skeleton: mechanical functions of living wood.

Wood is well defined as an engineering material. However, living wood in the tree is often regarded only as a passive skeleton consisting of a sophisticated pipe system for the ascent of sap and a tree-like structure made of a complex material to resist external forces. There are two other active key roles of living wood in the field of biomechanics: (i) additive manufacturing of the whole structure by cell division and expansion, and (ii) a 'muscle' function of living fibres or tracheids generating forces at the sapwood periphery. The living skeleton representing most of the sapwood is a mere accumulation of dead tracheids and libriform fibres after their programmed cell death. It keeps a record of the two active roles of living wood in its structure, chemical composition, and state of residual stresses. Models and field experiments define four biomechanical traits based on stem geometry and parameters of wood properties resulting from additive manufacturing and force generation. Geometric parameters resulting from primary and secondary growth play the larger role. Passive wood properties are only secondary parameters, while dissymmetric force generation is key for movement, posture control, and tree reshaping after accidents.