Edges impose planar alignment in nematic monolayers by directing cell elongation and enhancing migration

Abstract

Boundaries play an important role in the emergence of nematic order in classical liquid crystal systems; we explore their importance in adhesive cells that form active nematics. In particular, we study how cells are affected by an edge, which in our experiments is a boundary between adhesive and non-adhesive domains on a planar surface. We find that such edges induce elongation and direct the migration of isolated fibroblasts. In confluent monolayers, these elongated cells co-align and migrate to form an active, two-dimensional nematic structure in which edges enforce planar alignment and provide local slip to streams of cells that move along them. On an adhesive square island of dimensions 1 mm × 1 mm, cells near the edges in confluent nematic monolayers have enhanced alignment and velocity. The corners of the adhesive island seed defects with signs that depend on the direction of the motion of the streams of cells that meet there. Distortions emerge with rotations of −π/2 to form a −1/4 defect for streams that move clockwise or counterclockwise, and +π/2 to form a +1/4 defect for converging streams. We explore how cells transmit alignment information to each other in the absence of an edge by studying cell pairs and find that while such pairs do co-align, this alignment is only transient and short lived. These results shed light on the importance of edges in imposing nematic order in confluent monolayers and how edges can be used as tools to pattern the long-range organization of cells for tissue engineering applications.