Abstract:
In many biological systems, microorganisms swim through complex polymeric fluids, and usually deform the medium at a rate faster than the inverse fluid relaxation time. We address the basic properties of such life at high Deborah number analytically by considering the small-amplitude swimming of a body in an arbitrary complex fluid. Using asymptotic analysis and differential geometry, we show that for a given swimming gait, the time-averaged leading-order swimming kinematics of the body can be expressed as an integral equation on the solution to a series of simpler Newtonian problems. We then use our results to demonstrate that Purcell's scallop theorem, which states that time-reversible body motion cannot be used for locomotion in a Newtonian fluid, breaks down in polymeric fluid environments.

Abstract:
We consider pressure-driven flows of electrolyte solutions in small channels or capillaries in which tracer particles are used to probe velocity profiles. Under the assumption that the double layer is thin compared to the channel dimensions, we show that the flow-induced streaming electric field can create an apparent slip velocity for the motion of the particles, even if the flow velocity still satisfies the no-slip boundary condition. In this case, tracking of particle would lead to the wrong conclusion that the no-slip boundary condition is violated. We evaluate the apparent slip length, compare with experiments, and discuss the implications of these results.

Abstract:
The biological fluids encountered by self-propelled cells display complex microstructures and rheology. We consider here the general problem of low-Reynolds number locomotion in a complex fluid. {Building on classical work on the transport of particles in viscoelastic fluids,} we demonstrate how to mathematically derive three integral theorems relating the arbitrary motion of an isolated organism to its swimming kinematics {in a non-Newtonian fluid}. These theorems correspond to three situations of interest, namely (1) squirming motion in a linear viscoelastic fluid, (2) arbitrary surface deformation in a weakly non-Newtonian fluid, and (3) small-amplitude deformation in an arbitrarily non-Newtonian fluid. Our final results, valid for a wide-class of {swimmer geometry,} surface kinematics and constitutive models, at most require mathematical knowledge of a series of Newtonian flow problems, and will be useful to quantity the locomotion of biological and synthetic swimmers in complex environments.

Abstract:
Actuating periodically an elastic filament in a viscous liquid generally breaks the constraints of Purcell's scallop theorem, resulting in the generation of a net propulsive force. This observation suggests a method to design simple swimming devices - which we call "elastic swimmers" - where the actuation mechanism is embedded in a solid body and the resulting swimmer is free to move. In this paper, we study theoretically the kinematics of elastic swimming. After discussing the basic physical picture of the phenomenon and the expected scaling relationships, we derive analytically the elastic swimming velocities in the limit of small actuation amplitude. The emphasis is on the coupling between the two unknowns of the problems - namely the shape of the elastic filament and the swimming kinematics - which have to be solved simultaneously. We then compute the performance of the resulting swimming device, and its dependance on geometry. The optimal actuation frequency and body shapes are derived and a discussion of filament shapes and internal torques is presented. Swimming using multiple elastic filaments is discussed, and simple strategies are presented which result in straight swimming trajectories. Finally, we compare the performance of elastic swimming with that of swimming microorganisms.

Abstract:
In these lecture notes, I will briefly review the fundamental physical principles of locomotion in fluids, with a particular emphasis on the low-Reynolds number world.

Abstract:
Purcell's scallop theorem states that swimmers deforming their shapes in a time-reversible manner ("reciprocal" motion) cannot swim. Using numerical simulations and theoretical calculations we show here that in a fluctuating environment, reciprocal swimmers undergo, on time scales larger than that of their rotational diffusion, diffusive dynamics with enhanced diffusivities, possibly by orders of magnitude, above normal translational diffusion. Reciprocal actuation does therefore lead to a significant advantage over non-motile behavior for small organisms such as marine bacteria.

Abstract:
Locomotion on small scales is dominated by the effects of viscous forces and, as a result, is subject to strong physical and mathematical constraints. Following Purcell's statement of the scallop theorem which delimitates the types of swimmer designs which are not effective on small scales, we review the different ways the constraints of the theorem can be escaped for locomotion purposes.

Abstract:
Flagella beating in complex fluids are significantly influenced by viscoelastic stresses. Relevant examples include the ciliary transport of respiratory airway mucus and the motion of spermatozoa in the mucus-filled female reproductive tract. We consider the simplest model of such propulsion and transport in a complex fluid, a waving sheet of small amplitude free to move in a polymeric fluid with a single relaxation time. We show that, compared to self-propulsion in a Newtonian fluid occurring at a velocity U_N, the sheet swims (or transports fluid) with velocity U / U_N = [1+De^2 (eta_s)/(eta) ]/[1+De^2], where eta_s is the viscosity of the Newtonian solvent, eta is the zero-shear-rate viscosity of the polymeric fluid, and De is the Deborah number for the wave motion, product of the wave frequency by the fluid relaxation time. Similar expressions are derived for the rate of work of the sheet and the mechanical efficiency of the motion. These results are shown to be independent of the particular nonlinear constitutive equations chosen for the fluid, and are valid for both waves of tangential and normal motion. The generalization to more than one relaxation time is also provided. In stark contrast with the Newtonian case, these calculations suggest that transport and locomotion in a non-Newtonian fluid can be conveniently tuned without having to modify the waving gait of the sheet but instead by passively modulating the material properties of the liquid.

Abstract:
Purcell's scallop theorem defines the type of motions of a solid body - reciprocal motions - which cannot propel the body in a viscous fluid with zero Reynolds number. For example, the flapping of a wing is reciprocal and, as was recently shown, can lead to directed motion only if its frequency Reynolds number, Re_f, is above a critical value of order one. Using elementary examples, we show the existence of oscillatory reciprocal motions which are effective for all arbitrarily small values of the frequency Reynolds number and induce net velocities scaling as (Re_f)^\alpha (alpha > 0). This demonstrates a continuous breakdown of the scallop theorem with inertia.

Abstract:
Bacteria predate plants and animals by billions of years. Today, they are the world's smallest cells yet they represent the bulk of the world's biomass, and the main reservoir of nutrients for higher organisms. Most bacteria can move on their own, and the majority of motile bacteria are able to swim in viscous fluids using slender helical appendages called flagella. Low-Reynolds-number hydrodynamics is at the heart of the ability of flagella to generate propulsion at the micron scale. In fact, fluid dynamic forces impact many aspects of bacteriology, ranging from the ability of cells to reorient and search their surroundings to their interactions within mechanically and chemically-complex environments. Using hydrodynamics as an organizing framework, we review the biomechanics of bacterial motility and look ahead to future challenges.