We show in this study that Salmonella cells, which do not upregulate flagellar gene expression during swarming, also do not increase flagellar numbers per m of cell length as determined by systematic counting of both flagellar filaments and hooks. Instead, doubling of the average length of a swarmer cell by suppression of cell division effectively doubles the number of flagella per cell. The highest agar concentration at which Salmonella cells swarmed increased from the normal 0.5% to 1%, either when flagella were overproduced or when expression of the FliL protein was enhanced in conjunction with stator proteins MotAB. We surmise that bacteria use the resulting increase in motor power to overcome the higher friction associated with harder agar. Higher flagellar numbers also suppress the swarming defect of mutants with changes in the chemotaxis pathway that were previously shown to be defective in hydrating their colonies. Here we show that the swarming defect of these mutants can also be suppressed by application of osmolytes to the surface of swarm agar. The "dry" colony morphology displayed by che mutants was also observed with other mutants that do not actively rotate their flagella. The flagellum/motor thus participates in two functions critical for swarming, enabling hydration and overriding surface friction. We consider some ideas for how the flagellum might help attract water to the agar surface, where there is no free water.
Swarming bacteria may be divided into two categories: robust swarmers, which can navigate across a hard agar surface (1.5% agar and above), and temperate swarmers, which can swarm only on a softer agar surface (0.5 to 0.8% agar) (1-3). Robust swarmers include polarly flagellated bacteria, which induce peritrichous flagellation upon surface contact, such as Azospirillum, Rhodospirillum, and Vibrio species, as well as the peritrichously flagellated Proteus species (4-6). These bacteria display a hyperflagellated and hyperelongated swarm cell morphology, which is dramatically different from their broth-grown (swimming) counterparts. Transcriptome studies have shown a swarming-specific gene expression program in the robust swarmers (7-10). In polarly flagellated bacteria, frictional forces on the surface are surmised to slow flagellar rotation and signal altered gene expression (11,12), an observation solidified by experiments demonstrating a direct relationship between motor speed and lateral flagellar (laf) gene transcription (13). How motor speed might be sensed and transduced to activate laf expression is unknown. This phenomenon is apparently unique to polarly flagellated bacteria.Temperate swarmers include Escherichia coli and Bacillus, Pseudomonas, Rhizobium, Salmonella, Serratia, and Yersinia species. Among these, Bacillus subtilis displays increased flagellar numbers and cell length (14), but this morphology is not as dramatic as that seen in the robust swarmers. In a Bacillus cereus transcriptome, several genes, including flagellar genes, were seen to be differentially regulat...