Single Nanoparticle Tracking Reveals Efficient Long-Distance Undercurrent Transport above Swarming Bacteria

Abstract

Flagellated bacteria move collectively in a swirling pattern on agar surfaces immersed in a thin layer of viscous "swarm fluid", but the role of this fluid in mediating the cooperation of the bacterial population is not well understood. Herein, we use gold nanorods (AuNRs) as single particle tracers to explore the spatiotemporal structure of the swarm fluid. We observed that individual AuNRs are transported in a plane of ~2 µm above the motile cells. They can travel for long distances (~800 μm) in a 2D plane at high speed without interferences from bacterial movements. The particles are apparently lifted up and transported by collective mixing of the small vortices around bacteria during localized clustering and de-clustering of the motile cells, exhibiting superdiffusive and non-Gaussian characteristics with alternating large-step jumps and confined lingering. Their motions are consistent with the Lévy walk (LW) model, revealing efficient transport flows above swarms. These flows provide obstacle-free highways for long-range material transportations, shed light on how swarming bacteria perform population-level communications, and reveal the essential role of the fluid phase on the emergence of large-scale synergy. This approach is promising for probing complex fluid dynamics and transports in other collective systems.