Anomalous elasticity of a cellular tissue vertex model.

Vertex models, such as those used to describe cellular tissue, have an energy controlled by deviations of each cell area and perimeter from target values. The constrained nonlinear relation between area and perimeter leads to new mechanical response. Here we provide a mean-field treatment of a highly simplified model: a uniform network of regular polygons with no topological rearrangements. Since all polygons deform in the same way, we only need to analyze the ground states and the response to deformations of a single polygon (cell). The model exhibits the known transition between a fluid/compatible state, where the cell can accommodate both target area and perimeter, and a rigid/incompatible state. We calculate and measure the mechanical resistance to various deformation protocols and discover that at the onset of rigidity, where a single zero-energy ground state exists, linear elasticity fails to describe the mechanical response to even infinitesimal deformations. In particular, we identify a breakdown of reciprocity expressed via different moduli for compressive and tensile loads, implying nonanalyticity of the energy functional. We give a pictorial representation in configuration space that reveals that the complex elastic response of the vertex model arises from the presence of two distinct sets of reference states (associated with target area and target perimeter). Our results on the critically compatible tissue provide a new route for the design of mechanical metamaterials that violate or extend classical elasticity.