Cluster and conquer: the morphodynamics of invasion of a compliant substrate by active rods

The colonisation of a soft passive material by motile cells such as bacteria is common in biology. The resulting colonies of the invading cells are often observed to exhibit intricate patterns whose morphology and dynamics can depend on a number of factors, particularly the mechanical properties of the substrate and the motility of the individual cells. We use simulations of a minimal 2D model of self-propelled rods moving through a passive compliant medium consisting of particles that offer elastic resistance before being plastically displaced from their equilibrium positions. It is observed that the clustering of active (self-propelled) particles is crucial for understanding the morphodynamics of colonisation. Clustering enables motile colonies to spread faster than they would have as isolated particles. The colonisation rate depends non-monotonically on substrate stiffness with a distinct maximum at a non-zero value of substrate stiffness. This is observed to be due to a change in the morphology of clusters. Furrow networks created by the active particles have a fractal-like structure whose dimension varies systematically with substrate stiffness but is less sensitive to particle activity. The power-law growth exponent of the furrowed area is smaller than unity, suggesting that, to sustain such extensive furrow networks, colonies must regulate their overall growth rate. Active rod-shaped cells such as bacteria colonize soft plastic materials by forming furrow networks. Rafts and trains of advancing rods enhance the speed of colonization. This enhancement and the furrow network morphology depend strongly on substrate stiffness.