Multipili attachment of bacteria with helixlike pili exposed to stress

A number of biomechanical properties of various types of pili expressed by Escherichia coli, predominantly their force-versus-elongation behavior, have previously been assessed in detail on a single pilus level. In vivo, however, bacteria bind in general to host cells by a multitude of pili, which presumably provides them with adhesion properties that differs from those of single pili. Based upon the previously assessed biomechanical properties of individual pili, this work presents a theoretical analysis of the adhesion properties of multipili-attaching bacteria expressing helixlike pili exposed to an external force. Expressions for the adhesion lifetime of dual- and multipili-attaching bacteria are derived and their validity is verified by Monte Carlo simulations. It is demonstrated that the adhesion lifetime of a multipili-binding bacterium depends to a large degree on the cooperativity of the attaching pili, which, in turn, depends strongly on their internal biomechanical properties, in particular their helixlike structure and its ability to elongate, which, in turn, depends on the intrinsic properties of the bonds, e.g., their lengths and activation energies. It is shown, for example, that a decrease in the length of a layer-to-layer bond in the rod of P pili, expressed by E. coli, by 50% leads to a decrease in the adhesion lifetime of a bacterium attaching by ten pili and exposed to a force of 500 pN by three orders of magnitude. The results indicate moreover that the intrinsic properties of the rod for this particular type of pili are optimized for multipili attachment under a broad range of external forces and presumably also to its in vivo environment. For example, P pili seems to be optimized to withstand a force exposure during approximately 3 s, which correspond to the time it takes for a bolus to pass a bacterium attached to the ureteral wall. Even though the results presented in this work apply quantitatively to one type of pilus, they are assumed to apply qualitatively to all helixlike pili systems expressing slip bonds.