Formation, collective motion, and merging of macroscopic bacterial aggregates

Chemotactic bacteria form emergent spatial patterns of variable cell density within cultures that are initially spatially uniform. These patterns are the result of chemical gradients that are created from the directed movement and metabolic activity of billions of cells. A recent study on pattern formation in wild bacterial isolates has revealed unique collective behaviors of the bacteria Enterobacter cloacae. As in other bacterial species, Enterobacter cloacae form macroscopic aggregates. Once formed, these bacterial clusters can migrate several millimeters, sometimes resulting in the merging of two or more clusters. To better understand these phenomena, we examine the formation and dynamics of thousands of bacterial clusters that form within a 22 cm square culture dish filled with soft agar over two days. At the macroscale, the aggregates display spatial order at short length scales, and the migration of cell clusters is superdiffusive, with a merging acceleration that is correlated with aggregate size. At the microscale, aggregates are composed of immotile cells surrounded by low density regions of motile cells. The collective movement of the aggregates is the result of an asymmetric flux of bacteria at the boundary. An agent-based model is developed to examine how these phenomena are the result of both chemotactic movement and a change in motility at high cell density. These results identify and characterize a new mechanism for collective bacterial motility driven by a transient, density-dependent change in motility.Author summary: Bacteria growing and swimming in soft agar often aggregate to form elaborate spatial patterns. Here we examine the patterns formed by the bacteria Enterobacter cloacae. An unusual behavior of these bacteria is the collective movement of cells after the initial aggregation into a tiny spot. Despite the majority of the cells within an aggregate being immotile at any point in the time, the flux of cells entering and leaving the aggregate, as motility is lost and regained in individual cells, led to a net, collective movement of the aggregate. These spots sometimes run into each other and combine. By looking at the cells within these spots under a microscope, we find that cells within each spot stop swimming. The process of switching back and forth between swimming and not swimming causes the movement and fusion of the spots. A numerical simulation shows that the migration and merging of these spots can be expected if the cells swim towards regions of space with high concentrations of attractant molecules and stop swimming in locations crowded with many cells. This work identifies a novel process through which populations of bacteria cooperate and control the movement of large groups of cells.