Epithelia Realize Nematopolar Topological Defect Structures

Abstract

We introduce a shape-based polar order parameter that captures the structural asymmetry of cells within epithelial monolayers. By combining bright-field imaging and traction force microscopy, we demonstrate that shape polarity serves as a unifying biomechanical metric, integrating the physical information encoded by nematic directors, principal stresses, and cellular motion. Furthermore, we show that the tissue organizes into a mixed polar-nematic phase, characterized by the coexistence of integer ($\pm 1$) and half-integer ($\pm 1/2$) defects. Through mechanical perturbations, we demonstrate that both substrate stiffness and cell-cell adhesion modulate the density of these excitations and the length of domain walls binding like-signed positive half-integer defects. Using a minimal continuum model of polar-nematic active matter, we establish that this mixed phase is fundamentally driven by the interplay of active stresses and polar-nematic elasticity. These findings provide a direct experimental evidence that epithelial monolayers behave as nematopolar matter, in which coupled polar and nematic elastic interactions jointly shape the active state