Self-patterning in bacterial biofilms: A mechanical blueprint.

Like other developmental systems, bacteria in biofilms reproducibly self-organize into ordered patterns during development. Despite recent progress in single-cell imaging, the physical principles underlying the long-range ordering in biofilm remain unknown. Using Vibrio cholerae as a model biofilm former, we uncover the mechanical blueprint that governs biofilm self-patterning. We show that, through mechanical instabilities at the single-cell level, a biofilm spontaneously creates spatially segregated zones of anisotropic growth, which drive macroscopic ordering. We develop a two-phase active nematic model to quantitatively predict the biofilm developmental process and to provide guidance for strategies to pattern biofilms. We verify our theory by mutagenesis and by optically manipulating cell growth. Our conclusions are potentially generalizable to a wide variety of developmental systems with growth-induced flows.