Many bacteria move through liquids and across surfaces by using flagella-filaments propelled by a membrane-embedded rotary motor. Much is known about the flagellum: its basic structure, the function of its individual motor components, and the regulation of its synthesis. However, we are only beginning to identify the dynamics of flagellar proteins and to understand how the motor structurally adapts to environmental stimuli. In this review, we discuss the external and cellular factors that influence the dynamics of stator complexes (the ion-conducting channels of the flagellar motor). We focus on recent discoveries suggesting that stator dynamics are a means for controlling flagellar function in response to different environments.KEYWORDS flagella, stators, MotAB, swarming, swimming, c-di-GMP T he flagellar motor is an intricate machine that drives the rotation of the flagellum by converting ion motive force to mechanical energy (1). Torque is generated via association between two main motor components: membrane-embedded stator complexes and the cytoplasmic rotor ( Fig. 1). On the basis of work with the model organism Escherichia coli, each stator complex is composed of protein subunits, MotA and MotB, assembled in a 4MotA-2MotB stoichiometry (2). MotB has been shown to interact with the P-ring protein FlgI (3) and is also thought to associate with peptidoglycan through its peptidoglycan-binding motif. The interactions made by the periplasmic domain of MotB likely serve to anchor stator complexes around the rotor, where the stators act as ion channels. It is thought that the ions passing through these channels are bound by a conserved aspartic acid residue in the transmembrane segment of MotB, causing conformational changes in MotA (4). These conformational changes are coupled to rotation by specific electrostatic interactions between MotA and the rotor protein FliG (5, 6). Work with Vibrio alginolyticus suggests that FliG contributes to both rotation and stator assembly (7).While stators are named for their "stationary" role as the nonrotating motor component, the composition of stators surrounding the motor is highly dynamic (Fig. 2). Stator dynamics were first demonstrated in "resurrection experiments." These experiments showed that stators could incorporate themselves into and restore rotation to paralyzed flagellar motors, with each successive stator incorporation resulting in a stepwise increase in motor speed (8, 9). Importantly, stochastic fluctuations in torque were also observed in these experiments, in which torque would periodically decrease in steps (8). Specifically, 11 distinct rotational speeds were observed in E. coli, indicating that a maximum of at least 11 stator complexes can engage in the motor at any one time in this species (10). Stator turnover was directly observed and measured by total internal reflection fluorescence microscopy combined with fluorescence recovery after photobleaching (FRAP) and fluorescence loss in photobleaching. These experiments demonstrated that green fluo...