Growth phases of an active tissue: determinate, indeterminate, and proportionate

Abstract

Growth may cease at a target size or continue throughout life: the determinate and indeterminate phenotypes. We develop an active viscoelastic continuum model of a tissue growing along one axis, in which cell division and death generate active stresses. We find two asymptotic states: one in which the tissue reaches a relative size fixed by its material parameters, and one in which it elongates linearly without bound. Which state is realised is set by the ratio of active stress to elastic modulus. The transition originates in a bound on the elastic stress the tissue can support: a sufficiently large activity can never be balanced. In a tissue made of parts with different material properties, the growing phase settles into fixed length proportions, set by the mechanical impedances of the parts rather than inherited; matching impedances to initial lengths preserves the proportions the tissue began with. Determinate, indeterminate and proportionate growth thus appear as regimes of one continuum mechanical framework.