Spectral approaches to stress relaxation in epithelial monolayers.

We investigate the viscoelastic relaxation to equilibrium of an isolated disordered planar epithelium described using the cell vertex model. In its standard form, the model is formulated as coupled evolution equations for the locations of vertices of confluent polygonal cells. Exploiting the model's gradient-flow structure, we use singular-value decomposition to project modes of deformation of vertices onto modes of deformation of cells. Expressing the dynamics in terms of operators of a discrete calculus, we show how eigenmodes of discrete Laplacian operators (specified by constitutive assumptions related to dissipation and mechanical energy) provide a spatial basis for evolving fields, and demonstrate how the operators can incorporate approximations of conventional spatial derivatives. We relate the spectrum of relaxation times to the eigenvalues of the Laplacians, modified by corrections that account for the fact that the cell network (and therefore the Laplacians) evolve during relaxation to an equilibrium prestressed state, providing the monolayer with geometric stiffness. While dilational modes of the Laplacians capture rapid relaxation in some circumstances, showing diffusive dynamics, geometric stiffness is typically a dominant source of monolayer rigidity, as we illustrate for monolayers exposed to unsteady stretching deformations.