Emergence of active nematics in bacterial biofilms

Abstract

Growing tissue and bacterial colonies are active matter systems where cell divisions and cellular motion generate active stress. Although they operate in the non-equilibrium regime, these biological systems can form large-scale ordered structures such as nematically aligned cells, topological defects, and fingerings. Mechanical instabilities also play an essential role during growth by generating large structural folding. How active matter dynamics and mechanical instabilities together develop large-scale order in growing tissue is not well understood. Here, we use chain forming Bacillus subtilis, also known as a biofilm, to study the direct relation between active stress and nematic ordering. We find that a bacterial biofilm has intrinsic length scales above which series of mechanical instabilities occur. Localized stress and friction control both linear buckling and edge instabilities. Remarkably, these instabilities develop nematically aligned cellular structures and create pairs of motile and stationary topological defects. We also observe that stress distribution across the biofilm strongly depends on the defect dynamics which can further initiate the formation of sporulation sites by creating three-dimensional structures. By investigating the development of bacterial biofilms and their mechanical instabilities we are proposing a new type of active matter system which provides a unique platform to study the essential roles of nematics in growing biological tissue.