Bacteriophage is known to infect motile strains of enteric bacteria by adsorbing randomly along the length of a f lagellar filament and then injecting its DNA into the bacterial cell at the filament base. Here, we provide evidence for a ''nut and bolt'' model for translocation of phage along the filament: the tail fiber of fits the grooves formed by helical rows of f lagellin monomers, and active f lagellar rotation forces the phage to follow the grooves as a nut follows the threads of a bolt.Bacteriophage infects motile strains of the genera Escherichia, Salmonella, and Serratia (1-3). Schade et al. (4) demonstrated translocation of phage along flagellar filaments by electron microscopy: phage heads tended to be full when attached to a filament but empty when attached to its base. When it was realized that flagellar filaments rotate, the suggestion was made that a phage moves along the filament ''like a nut on a bolt'' (5). This observation was consistent with the finding that mutants with straight filaments, while nonmotile, remain fully sensitive to (6). Ravid and Eisenbach (7) found that the number of phage particles adsorbed by cells, i.e., removed from the supernatant fraction, correlated only with the fraction of the population of cells whose flagella rotated incessantly; it did not correlate with the direction of rotation. This finding was not consistent with a nut and bolt model. Yamaguchi et al. (8) found conditions in which unflagellated bacteria are sensitive to -phage: drops of concentrated phage cleared spots on the surfaces of hard agar plates. However, this sensitivity might have resulted simply from the large multiplicity of phage. Attempts to visualize phage translocation by dark-field or differential-interference-contrast microscopy have not been successful: one can see -phage attached to filaments but not their travel in either direction (R.M. Macnab, personal communication; ref. 9). However, these experiments required low flagellar rotation rates. The prevailing view seems to be that phage particles adsorb and desorb, stepping along the filament in a one-dimensional random walk.If flagellar rotation drives translocation in the manner of a nut on a bolt, then a bacterial strain's sensitivity to -phage has three mechanical requirements: flagellar rotation, the correct direction of flagellar rotation, and the correct pattern of grooves on the surface of the flagellar filament. By measuring infectivity directly, we asked whether these mechanical constraints are determinants of sensitivity to -phage.