Free surface flows driven by boundary undulations are observed in many biological phenomena, including the feeding and locomotion of water snails. To simulate the feeding strategy of apple snails, we develop a centimetric robotic undulator that drives a thin viscous film of liquid with the wave speed $V_w$. Our experimental results demonstrate that the behaviour of the net fluid flux $Q$ strongly depends on the Reynolds number $Re$. Specifically, in the limit of vanishing $Re$, we observe that $Q$ varies non-monotonically with $V_w$, which has been successfully rationalised by Pandey et al. (Nat. Commun., vol. 14, no. 1, 2023, p. 7735) with the lubrication model. By contrast, in the regime of finite inertia (${Re} \sim O(1)$), the fluid flux continues to increase with $V_w$ and completely deviates from the prediction of lubrication theory. To explain the inertia-enhanced pumping rate, we build a thin-film, two-dimensional model via the asymptotic expansion in which we linearise the effects of inertia. Our model results match the experimental data with no fitting parameters and also show the connection to the corresponding free surface shapes $h_2$. Going beyond the experimental data, we derive analytical expressions of $Q$ and $h_2$, which allow us to decouple the effects of inertia, gravity, viscosity and surface tension on free surface pumping over a wide range of parameter space.

Immersed nonlinear elements are prevalent in biological systems that require a preferential flow direction, such as the venous and the lymphatic system. We investigate here a certain class of models where the fluid is driven by peristaltic pumping and the nonlinear elements are ideal valves that completely suppress backflow. This highly nonlinear system produces discontinuous solutions that are difficult to study. We show that, as the density of valves increases, the pressure and flow are well approximated by a continuum of valves which can be analytically treated, and we demonstrate through numeric simulation that the approximation works well even for intermediate valve densities. We find that the induced flow is linear in the peristaltic amplitude for small peristaltic forces and, in the case of sinusoidal peristalsis, is independent of pumping direction. Despite the continuum approximation used, the physical valve density is accounted for by modifying the resistance of the fluid appropriately. The suppression of backflow causes a net benefit in adding valves when the valve density is low, but once the density is high enough, valves predominately suppress forward flow, suggesting there is an optimum number of valves per wavelength. The continuum model for peristaltic pumping through an array of valves presented in this work can eventually provide insights about the design and operating principles of complex flow networks with a broad class of nonlinear elements.

A theoretical analysis is presented of peristaltic pumping down a narrow conduit with permeable walls, motivated by the flushing action of lugworms and other marine organisms in sandy burrows. Flow in the conduit is dealt with using lubrication theory; the leakage into the surrounding medium is taken into account by exploiting slender-body theory to solve the associated Darcy problem. By adopting a model for the local force balance on the pumping surface, we bridge between the limits in which the pump operates with either fixed load or displacement. In the latter limit we characterize peristaltic waves with either fixed form or ones that partially collapse the conduit. We construct pump characteristics (the relation between the mean flux and net pressure drop) when the burrow wall is impermeable and pressures are fixed at each end, and compare the results with existing laboratory experiments performed on lugworms. We then consider how the peristaltic dynamics is changed when the wall is made permeable. Last, we consider pumping along an impermeable burrow into a leaky head shaft. The results reveal that the permeability of the conduit wall or end can greatly impact the direction and strength of the recirculating flow.

In this study, for the first time, we consider longitudinal motion of the walls during peristalsis in a distensible tube and how this affects backward (or retrograde) flow, i.e. peristaltic reflux. Building on the analytical model developed by Shapiro et al. (J. Fluid Mech., vol. 37, no. 4, 1969, pp. 799–825) based on lubrication theory, we model peristalsis as a two-dimensional infinite sinusoidal wavetrain. We develop an objective function with high mechanical pumping efficiency and low reflux to find optimal peristalsis conditions. We show that optimal wall longitudinal motion contributes substantially to limiting reflux during peristalsis. The results suggest that the optimal form of wall longitudinal velocity is a linear function of the wall transverse coordinate, moving forward with the wave when the tube is distended and retracting when contracted. Our results are in general agreement with clinical observations of ureteral peristalsis.

This paper addresses peristaltic flow induced in a non-axisymmetric annular tube by a periodic small-amplitude wave of arbitrary shape propagating axially along its inner surface, assumed to be a circular cylinder. The study is motivated by recent in vivo experimental observations pertaining to the flow of cerebrospinal fluid along the perivascular spaces of cerebral arteries. The analysis employs the lubrication approximation, describing low-Reynolds-number peristaltic flow in the long-wavelength approximation. Closed-form analytic expressions are derived for the average pumping rate in infinitely long tubes and also in tubes of finite length. Consideration is also given to the transverse motion arising in non-axisymmetric tubes. For small-amplitude waves, the solution is reduced to the integration of a parameter-free Stokes-flow problem, which is solved for relevant cross-sectional shapes, with closed-form analytical results derived for thin canals.

Using numerical simulations, we probe the fluid flow in an axisymmetric peristaltic vessel fitted with elastic bi-leaflet valves. In this biomimetic system that mimics the flow generated in lymphatic vessels, we investigate the effects of the valve and vessel properties on pumping performance of the valved peristaltic vessel. The results indicate that valves significantly increase pumping by reducing backflow. The presence of valves, however, increases the viscous resistance, therefore requiring greater work compared to valveless vessels. The benefit of the valves is the most significant when the fluid is pumped against an adverse pressure gradient and for low vessel contraction wave speeds. We identify the optimum vessel and valve parameters leading to the maximum pumping efficiency. We show that the optimum valve elasticity maximizes the pumping flow rate by allowing the valve to block the backflow more effectively while maintaining low resistance during the forward flow. We also examine the pumping in vessels where the vessel contraction amplitude is a function of the adverse pressure gradient, as found in lymphatic vessels. We find that, in this case, the flow is limited by the work generated by the contracting vessel, suggesting that the pumping in lymphatic vessels is constrained by the performance of the lymphatic muscle. Given the regional heterogeneity of valve morphology observed throughout the lymphatic vasculature, these results provide insight into how these variations might facilitate efficient lymphatic transport in the vessel's local physiologic context.

The two-dimensional laminar flow of a viscous fluid induced by peristalsis due to a moving wall wave has been studied previously for a rectangular channel, a circular tube and a concentric circular annulus. Here, we study peristaltic flow in a non-axisymmetric annular tube: in this case, the flow is three-dimensional, with motions in the azimuthal direction. This type of geometry is motivated by experimental observations of the pulsatile flow of cerebrospinal fluid along perivascular spaces surrounding arteries in the brain, which is at least partially driven by peristaltic pumping due to pulsations of the artery. These annular perivascular spaces are often eccentric and the outer boundary is seldom circular: their cross-sections can be well matched by a simple, adjustable model consisting of an inner circle (the outer wall of the artery) and an outer ellipse (the outer edge of the perivascular space), not necessarily concentric. We use this geometric model as a basis for numerical simulations of peristaltic flow: the adjustability of the model makes it suitable for other applications. We concentrate on the general effects of the non-axisymmetric configuration on the flow and do not attempt to specifically model perivascular pumping. We use a finite-element scheme to compute the flow in the annulus driven by a propagating sinusoidal radial displacement of the inner wall. Unlike the peristaltic flow in a concentric circular annulus, the flow is fully three-dimensional: azimuthal pressure variations drive an oscillatory flow in and out of the narrower gaps, inducing an azimuthal wiggle in the streamlines. We examine the dependence of the flow on the elongation of the outer elliptical wall and the eccentricity of the configuration. We find that the time-averaged volumetric flow is always in the same direction as the peristaltic wave and decreases with increasing ellipticity or eccentricity. The additional shearing motion in the azimuthal direction will increase mixing and enhance Taylor dispersion in these flows, effects that might have practical applications.

A laboratory experiment shows that fluid can be pumped from one end to another in a narrow channel whose sawtooth walls vibrate transversely opposite to each other. The phenomenon is referred to as ratcheting fluid. Inspired by this, we put forward here a theory describing the rectified steady flow, and the net directional pumping. In a conformally transformed plane, the induced steady streaming in the Stokes boundary layer of the oscillatory flow is analysed, elucidating the driving mechanism that is due to the nonlinearity and viscosity. The solution of the stream function is given, showing the complex spatial structure of the induced steady flow and its spatial average that is related to the directional pumping. Whereas the wall sawtooth shape is the primary source of asymmetry, the difference in entrance and exit flow conditions due to the geometries at channel ends is found to be a secondary source to break the left–right symmetry of the system. This latter can affect the net directional transport of fluid, in particular in short channels with a small number of sawteeth. Various influences on the net pumping rate are analysed.

We examine the fluid mechanics of drinking in nature. We classify the drinking strategies of a broad range of creatures according to the principal forces involved, and present physical pictures for each style. Simple scaling arguments are developed and tested against existing data. While suction is the most common drinking strategy, various alternative styles have evolved among creatures whose morphological, physiological and environmental constraints preclude it. Particular attention is given to creatures small relative to the capillary length, whose drinking styles rely on relatively subtle interfacial effects. We also discuss attempts to rationalize various drinking strategies through consideration of constrained optimization problems. Some biomimetic applications are discussed.

A model is derived for long peristaltic waves propagating steadily down a fluid-filled, axisymmetric tube. The waves are driven by imposing a radial force of prescribed form on the tube. The resulting deformation of the tube wall is modelled using linear elasticity and the internal flow using the lubrication approximation. Numerical solutions for periodic wave trains and solitary waves are presented, along with asymptotic solutions at both small and large forcing amplitudes. Large-amplitude periodic waves are characterized by narrow blisters adjoining long occluded sections of the tube, whereas a solitary wave of strong contraction produces a long inflated bow wave that propels a large quantity of fluid. A measure of pumping efficacy is given by the ratio of the net fluid displacement to the power input, and is highest for a large-amplitude solitary wave.

A mathematical model is developed for long peristaltic waves propelling a suspended rigid object down a fluid-filled axisymmetric tube. The fluid flow is described using lubrication theory and the deformation of the tube using linear elasticity. The object is taken to be either an infinitely long rod of constant radius or a parabolic-shaped lozenge of finite length. The system is driven by a radial force imposed on the tube wall that translates at constant speed down the tube axis and with a form chosen to generate a periodic wave train or a solitary wave. These waves exert a traction on the enclosed object, forcing it into motion. Periodic waves drive the infinite rod at a speed that attains a maximum at a moderate forcing amplitude and approaches approximately one quarter of the wave speed in the large-amplitude limit. The finite lozenge can be entrained and driven at the same speed as a solitary wave or periodic wave train if the forcing is sufficiently strong. For weaker forcing, the lozenge is either left behind the solitary wave or interacts repeatedly with the waves in the periodic train to generate stuttering forward progress. The threshold forcing amplitude for entrainment increases weakly with the radial span of the enclosed object, but strongly with the axial length, with entrainment becoming impossible if the object is too long.

Peristaltic pumping of fluid is a fundamental method of transport in many biologicalprocesses. In some instances, particles of appreciable size are transported along with thefluid, such as ovum transport in the oviduct or kidney stones in the ureter. In some ofthese biological settings, the fluid may be viscoelastic. In such a case, a nonlinearconstitutive equation to describe the evolution of the viscoelastic contribution to thestress tensor must be included in the governing equations. Here we use an immersedboundary framework to study peristaltic transport of a macroscopic solid particle in aviscoelastic fluid governed by a Navier-Stokes/Oldroyd-B model. Numerical simulations ofperistaltic pumping as a function of Weissenberg number are presented. We examine thespatial and temporal evolution of the polymer stress field, and also find that theviscoelasticity of the fluid does hamper the overall transport of the particle in thedirection of the wave.

Two types of wall actuation in channel flow are considered: travelling waves of wall deformation (peristalsis) and travelling waves of blowing and suction. The flow response and its mechanisms are analysed using nonlinear and weakly nonlinear computations. We show that both actuations induce a flux in the channel in the absence of an imposed pressure gradient and can thus be characterized as pumping. In the context of flow control, pumping and drag reduction are strongly connected, and we seek to define them properly based on these two actuation examples. Movies showing the flow motion for the two types of actuation are available with the online version of this paper (journals.cambridge.org/FLM).