Mehdi Jabbarzadeh

Helical and rod-shaped bacteria swim in helical trajectories with little additional propulsion from helical shape

Helical bacteria, like the stomach pathogen H. pylori, swim only about 15% faster than their isogenic rod-shaped mutants. It has frequently been hypothesized that the helical body shapes of flagellated bacteria may yield some advantage in swimming ability. In particular, the helical-shaped pathogen Helicobacter pylori is often claimed to swim like a corkscrew through its harsh gastric habitat, but there has been no direct confirmation or quantification of such claims. Using fast time-resolution and high-magnification two-dimensional (2D) phase-contrast microscopy to simultaneously image and track individual bacteria in bacterial broth as well as mucin solutions, we show that both helical and rod-shaped H. pylori rotated as they swam, producing a helical trajectory. Cell shape analysis enabled us to determine shape as well as the rotational and translational speed for both forward and reverse motions, thereby inferring flagellar kinematics. Using the method of regularized Stokeslets, we directly compare observed speeds and trajectories to numerical calculations for both helical and rod-shaped bacteria in mucin and broth to validate the numerical model. Although experimental observations are limited to select cases, the model allows quantification of the effects of body helicity, length, and diameter. We find that due to relatively slow body rotation rates, the helical shape makes at most a 15% contribution to propulsive thrust. The effect of body shape on swimming speeds is instead dominated by variations in translational drag required to move the cell body. Because helical cells are one of the strongest candidates for propulsion arising from the cell body, our results imply that quite generally, swimming speeds of flagellated bacteria can only be increased a little by body propulsion.

Choice of computational method for swimming and pumping with nonslender helical filaments at low Reynolds number

The flows induced by biological and artificial helical filaments are important to many possible applications including microscale swimming and pumping. Microscale helices can span a wide range of geometries, from thin bacterial flagella to thick helical bacterial cell bodies. While the proper choice of numerical method is critical for obtaining accurate results, there is little guidance about which method is optimal for a specified filament geometry. Here, using two physical scenarios — a swimmer with a head and a pump — we establish guidelines for the choice of numerical method based on helical radius, pitch, and filament thickness. For a range of helical geometries that encompass most natural and artificial helices, we create benchmark results using a surface distribution of regularized Stokeslets and then evaluate the accuracy of resistive force theory, slender body theory, and a centerline distribution of regularized Stokeslets. For the centerline distribution of regularized Stokeslets or slender body theory, we tabulate appropriate blob size and Stokeslet spacing or segment length, respectively, for each geometry studied. Finally, taking the computational cost of each method into account, we present the optimal choice of numerical method for each filament geometry as a guideline for future investigations involving filament-induced flows.

Bacteria-inspired nanorobots with flagellar polymorphic transformations and bundling

Wirelessly controlled nanoscale robots have the potential to be used for both in vitro and in vivo biomedical applications. So far, the vast majority of reported micro- and nanoscale swimmers have taken the approach of mimicking the rotary motion of helical bacterial flagella for propulsion, and are often composed of monolithic inorganic materials or photoactive polymers. However, currently no man-made soft nanohelix has the ability to rapidly reconfigure its geometry in response to multiple forms of environmental stimuli, which has the potential to enhance motility in tortuous heterogeneous biological environments. Here, we report magnetic actuation of self-assembled bacterial flagellar nanorobotic swimmers. Bacterial flagella change their helical form in response to environmental stimuli, leading to a difference in propulsion before and after the change in flagellar form. We experimentally and numerically characterize this response by studying the swimming of three flagellar forms. Also, we demonstrate the ability to steer these devices and induce flagellar bundling in multi-flagellated nanoswimmers.

Bipolar lophotrichous Helicobacter suis combine extended and wrapped flagella bundles to exhibit multiple modes of motility

The swimming strategies of unipolar flagellated bacteria are well known but little is known about how bipolar bacteria swim. Here we examine the motility of Helicobacter suis, a bipolar gastric-ulcer-causing bacterium that infects pigs and humans. Phase-contrast microscopy of unlabeled bacteria reveals flagella bundles in two conformations, extended away from the body (E) or flipped backwards and wrapped (W) around the body. We captured videos of the transition between these two states and observed three different swimming modes in broth: with one bundle rotating wrapped around the body and the other extended (EW), both extended (EE), and both wrapped (WW). Only EW and WW modes were seen in porcine gastric mucin. The EW mode displayed ballistic trajectories while the other two displayed superdiffusive random walk trajectories with slower swimming speeds. Separation into these two categories was also observed by tracking the mean square displacement of thousands of trajectories at lower magnification. Using the Method of Regularized Stokeslets we numerically calculate the swimming dynamics of these three different swimming modes and obtain good qualitative agreement with the measurements, including the decreased speed of the less frequent modes. Our results suggest that the extended bundle dominates the swimming dynamics.

Dynamic instability in the hook-flagellum system that triggers bacterial flicks

Dynamical bending, buckling, and polymorphic transformations of the flagellum are known to affect bacterial motility, but run-reverse-flick motility of monotrichous bacteria also involves the even more flexible hook connecting the flagellum to its rotary motor. Although flick initiation has been hypothesized to involve either static Euler buckling or dynamic bending of the hook, the precise mechanism of flick initiation remains unknown. Here, we find that flicks initiate via a dynamic instability requiring flexibility in both the hook and flagellum. We obtain accurate estimates of forces and torques on the hook that suggest that flicks occur for stresses below the (static) Euler buckling criterion, then provide a mechanistic model for flick initiation that requires combined bending of the hook and flagellum. We calculate the triggering torque-stiffness ratio and find that our predicted onset of dynamic instability corresponds well with experimental observations.