Dynamics of swimming bacteria: transition to directional order at high concentration

At high cell concentrations, bacterial suspensions are known to develop a state of collective swimming (the "zooming bionematic phase," or ZBN) characterized by transient, recurring regions of coordinated motion greatly exceeding the size of individual cells. Recent theoretical studies of semidilute suspensions have suggested that long-range hydrodynamic interactions between swimming cells are responsible for long-wavelength instabilities that lead to these patterns, while models appropriate for higher concentrations have suggested that steric interactions between elongated cells play an important role in the self-organization. Using particle imaging velocimetry in well-defined microgeometries, we examine the statistical properties of the transition to the ZBN in suspensions of Bacillus subtilis, with particular emphasis on the distribution of cell swimming speeds and its correlation with orientational order. This analysis reveals a nonmonotonic relationship between mean cell swimming speed and cell concentration, with a minimum occurring near the transition to the ZBN. Regions of high orientational order in the ZBN phase have locally high swimming speeds, while orientationally disordered regions have lower speeds. A model for steric interactions in concentrated suspensions and previous observations on the kinetics of flagellar rebundling associated with changes in swimming direction are used to explain this observation. The necessity of incorporating steric effects on cell swimming in theoretical models is emphasized.