Bacterial swarming is a type of motility characterized by a rapid and collective migration of bacteria on surfaces. Most swarming species form densely packed dynamic clusters in the form of whirls and jets, in which hundreds of rod-shaped rigid cells move in circular and straight patterns, respectively. Recent studies have suggested that short-range steric interactions may dominate hydrodynamic interactions and that geometrical factors, such as a cell's aspect ratio, play an important role in bacterial swarming. Typically, the aspect ratio for most swarming species is only up to 5, and a detailed understanding of the role of much larger aspect ratios remains an open challenge. Here we study the dynamics of Paenibacillus dendritiformis C morphotype, a very long, hyperflagellated, straight (rigid), rod-shaped bacterium with an aspect ratio of ϳ20. We find that instead of swarming in whirls and jets as observed in most species, including the shorter T morphotype of P. dendritiformis, the C morphotype moves in densely packed straight but thin long lines. Within these lines, all bacteria show periodic reversals, with a typical reversal time of 20 s, which is independent of their neighbors, the initial nutrient level, agar rigidity, surfactant addition, humidity level, temperature, nutrient chemotaxis, oxygen level, illumination intensity or gradient, and cell length. The evolutionary advantage of this unique back-and-forth surface translocation remains unclear.
Motile bacteria are able to colonize surfaces using various motility mechanisms (1). One efficient method includes flagellation-based cell motion in conjunction with collective lubrication (typically by secretion of surfactants) to enable fast expansion on hard surfaces. This mode of "bacterial swarming" that has been studied extensively for many species (1-15) enables rapid colony expansion (up to centimeters per hour). Swarming is often marked by hundreds of cells moving in a coordinated fashion while generating whirl and jet patterns.Studies of the collective dynamics of swarming have examined multiple aspects of motility. On the macroscopic level, it was discovered that swarming colonies show an advantage over liquid cultures in that they exhibit an increased resistance to antimicrobials (1,4,5,(15)(16)(17)(18)(19)(20). Studies of collective secretions of signaling and quorum-sensing molecules have shown how interactions between cells in swarming colonies are controlled (11) and exposed the identification of associated genetic manipulations and upregulated proteins that control biosurfactant secretions and flagellar behavior. On the single-cell level, attention was given to swarm cell trajectories and the ways in which these trajectories are determined by flagellar motion (8,(21)(22)(23). A combination of experiments (2, 3, 14, 15, 24-38) and theory (39-43) suggests that hydrodynamic interactions play a significant role in this social form of migration.Hydrodynamic interactions may not always be the dominant physical mechanism controlling bacterial motion....