Vesicles under a shear flow exhibit a tank-treading motion of their membrane, while their long axis points with an angle < 45 degrees with respect to the shear stress if the viscosity contrast between the interior and the exterior is not large enough. Above a certain viscosity contrast, the vesicle undergoes a tumbling bifurcation, a bifurcation which is known for red blood cells. We have recently presented the full numerical analysis of this transition. In this paper, we introduce an analytical model that has the advantage of being both simple enough and capturing the essential features found numerically. The model is based on general considerations and does not resort to the explicit computation of the full hydrodynamic field inside and outside the vesicle.
We add thermal noise consistently to reduced models of undeformable vesicles and capsules in shear flow and derive analytically the corresponding stochastic equations of motion. We calculate the steady-state probability distribution function and construct the corresponding phase diagrams for the different dynamical regimes. For fluid vesicles, we predict that at small shear rates thermal fluctuations induce a tumbling motion for any viscosity contrast. For elastic capsules, due to thermal mixing, an intermittent regime appears in regions where deterministic models predict only pure tank treadi…
An analytical theory is developed to describe the dynamics of a closed lipid bilayer membrane (vesicle) freely suspended in a general linear flow. Considering a nearly spherical shape, the solution to the creeping-flow equations is obtained as a regular perturbation expansion in the excess area. The analysis takes into account the membrane fluidity, incompressibility and resistance to bending. The constraint for a fixed total area leads to a non-linear shape evolution equation at leading order. As a result two regimes of vesicle behavior, tank-treading and tumbling, are predicted depending on …
Despite its significance in microfluidics, the effect of confinement on the transition from the tank-treading (steady motion) to the tumbling (unsteady motion) dynamical state of deformable micro-particles has not been studied in detail. In this paper, we investigate the dynamics of a single viscous vesicle under confining shear as a general model system for red blood cells, capsules, or viscous droplets. The transition from tank-treading to tumbling motion can be triggered by the ratio between internal and external fluid viscosities. Here, we show that the transition can be induced solely by …
Despite the recent upsurge of theoretical reduced models for vesicle shape dynamics, comparisons with experiments have not been accomplished. We review the implications of some of the recently proposed models for vesicle dynamics, especially the Tumbling-Trembling domain regions of the phase plane and show that they all fail to capture the essential behavior of real vesicles for excess areas, Δ, greater than 0.4. We emphasize new observations of shape harmonics and the role of thermal fluctuations.
We reveal that under moderate shear stress (of the order of 0.1 Pa) red blood cells present an oscillation of their inclination (swinging) superimposed to the long-observed steady tanktreading (TT) motion. A model based on a fluid ellipsoid surrounded by a visco-elastic membrane initially unstrained (shape memory) predicts all observed features of the motion: an increase of both swinging amplitude and period (1/2 the TT period) upon decreasing the shear stress, a shear stress-triggered transition towards a narrow shear stress-range intermittent regime of successive swinging and tumbling, and a…
We present a model for the dynamics of fluid vesicles in linear flow which consistently includes thermal fluctuations and nonlinear coupling between different modes. At the transition between tank-treading and tumbling, we predict a trembling motion which is at odds with the known deterministic motions and for which thermal noise is strongly amplified. In particular, highly asymmetric shapes are observed even though the deterministic flow only allows for axisymmetric ones. Our results explain quantitatively recent experimental observations [Levant and Steinberg, Phys. Rev. Lett. 109, 268103 (2…
Growing living cultures of Escherichia coli bacteria were investigated using real-time in situ rheology and rheo-imaging measurements. In the early stages of growth (lag phase), and when subjected to a constant stationary shear, the viscosity slowly increases with the cell's population. As the bacteria reach the exponential phase of growth, the viscosity increases rapidly, with sudden and temporary abrupt decreases and recoveries. At a certain stage, corresponding grossly to the late phase of growth, when the population stabilises, the viscosity also keeps its maximum constant value, with drop…
Vesicles and many biological membranes are made of two monolayers of lipid molecules and form closed lipid bilayers. The dynamical behaviour of vesicles is very complex and a variety of forms and shapes appear. Lipid bilayers can be considered as a surface fluid and hence the governing equations for the evolution include the surface (Navier--)Stokes equations, which in particular take the membrane viscosity into account. The evolution is driven by forces stemming from the curvature elasticity of the membrane. In addition, the surface fluid equations are coupled to bulk (Navier--)Stokes equatio…
The viscosity of lipid bilayer membranes plays an important role in determining the diffusion constant of embedded proteins and the dynamics of membrane deformations, yet it has historically proven very difficult to measure. Here we introduce a new method based on quantification of the large-scale circulation patterns induced inside vesicles adhered to a solid surface and subjected to simple shear flow in a microfluidic device. Particle Image Velocimetry based on spinning disk confocal imaging of tracer particles inside and outside of the vesicle, and tracking of phase-separated membrane domai…
The microscopic dynamics of objects suspended in a fluid determines the macroscopic rheology of a suspension. For example, as shown by Danker and Misbah [Phys. Rev. Lett. {\bf 98}, 088104 (2007)], the viscosity of a dilute suspension of fluid-filled vesicles is a non-monotonic function of the viscosity contrast (the ratio between the viscosities of the internal encapsulated and the external suspending fluids) and exhibits a minimum at the critical point of the tank-treading-to-tumbling transition. By performing numerical simulations, we recover this effect and demonstrate that it persists for …
The hydrodynamic interaction of two deformable vesicles in shear flow induces a net displacement, in most cases an increase of their distance in the transverse direction. The statistical average of these interactions leads to shear-induced diffusion in the suspension, both at the level of individual particles which experience a random walk made of successive interactions, and at the level of suspension where a non-linear down-gradient diffusion takes place, an important ingredient in the structuring of suspension flows. We make an experimental and computational study of the interaction of a pa…
Recent experiments have shown that when a near-hemispherical lipid vesicle attached to a solid surface is subjected to a simple shear flow it exhibits a pattern of membrane circulation much like a dipole vortex. This is in marked contrast to the toroidal circulation that would occur in the related problem of a drop of immiscible fluid attached to a surface and subjected to shear. This profound difference in flow patterns arises from the lateral incompressibility of the membrane, which restricts the observable flows to those in which the velocity field in the membrane is two-dimensionally diver…
We review the dynamical behavior of giant fluid vesicles in various types of external hydrodynamic flow. The interplay between stresses arising from membrane elasticity, hydrodynamic flows, and the ever present thermal fluctuations leads to a rich phenomenology. In linear flows with both rotational and elongational components, the properties of the tank-treading and tumbling motions are now well described by theoretical and numerical models. At the transition between these two regimes, strong shape deformations and amplification of thermal fluctuations generate a new regime called trembling. I…
Vesicles are involved in a vast variety of transport processes in living organisms. Additionally, they serve as a model for the dynamics of cell suspensions. Predicting the rheological properties of their suspensions is still an open question, as even the interaction of pairs is yet to be fully understood. Here we analyse the effect of a single vesicle, undergoing tank-treading motion, on its surrounding shear flow by studying the induced disturbance field $δ\vec{V}$, the difference between the velocity field in its presence and absence. The comparison between experiments and numerical simulat…
The distribution of Red Blood Cells in a confined channel flow is inhomogeneous and shows a marked depletion near the walls due to a competition between migration away from the walls and shear-induced diffusion resulting from interactions between particles. We investigated the lift of RBCs in a shear flow near a wall and measured a significant lift velocity despite the tumbling motion of cells. We also provide values for the collective and anisotropic shear-induced diffusion of a cloud of RBCs, both in the direction of shear and in the direction of vorticity. A generic down-gradient subdiffusi…
The dynamics of red blood cells (RBCs) in oscillatory shear flow was studied using differential equations of three variables: a shape parameter, the inclination angle $θ$, and phase angle $ϕ$ of the membrane rotation. In steady shear flow, three types of dynamics occur depending on the shear rate and viscosity ratio. i) tank-treading (TT): $ϕ$ rotates while the shape and $θ$ oscillate. ii) tumbling (TB): $θ$ rotates while the shape and $ϕ$ oscillate. iii) intermediate motion: both $ϕ$ and $θ$ rotate synchronously or intermittently. In oscillatory shear flow, RBCs show various dynamics based on…
von Willebrand Factor is a mechano-sensitive protein circulating in blood that mediates platelet adhesion to subendothelial collagen and platelet aggregation at high shear rates. Its hemostatic function and thrombogenic effect, as well as susceptibility to enzymatic cleavage, are regulated by a conformational change from a collapsed globular state to a stretched state. Therefore, it is essential to account for the conformation of the vWF multimers when modeling vWF-mediated thrombosis or vWF degradation. We introduce a continuum model of vWF unfolding that is developed within the framework of …
Red blood cells (RBCs) -- erythrocytes -- suspended in plasma tend to aggregate and form rouleaux. During aggregation the first stage consists in the formation of RBC doublets [Blood cells, molecules, and diseases 25, 339 (1999)]. While aggregates are normally dissociated by moderate flow stresses, under some pathological conditions the aggregation becomes irreversible, which leads to high blood viscosity and vessel occlusion. We perform here two-dimensional simulations to study the doublet dynamics under shear flow in different conditions and its impact on rheology. We sum up our results on t…
Inhaled nanoparticles (< 100 nm) reaching the deep lung region first interact with the pulmonary surfactant, a thin lipid film lining the alveolar epithelium. To date, most biophysical studies have focused on particle induced modifications of the film interfacial properties. In comparison, there is less work on the surfactant bulk properties, and on their changes upon particle exposure. Here we study the viscoelastic properties of a biomimetic pulmonary surfactant in the presence of various engineered nanoparticles. The microrheology technique used is based on the remote actuation of micron-si…
