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We develop a mean-field model to examine the stability of a `quasi-2D suspension' of elongated particles embedded within a viscous membrane. This geometry represents several biological and synthetic settings, and we reveal mechanisms by which the anisotropic mobility of particles interacts with long-ranged viscous membrane hydrodynamics. We first show that a system of slender rod-like particles driven by a constant force is unstable to perturbations in concentration -- much like sedimentation in analogous 3D suspensions -- so long as membrane viscous stresses dominate. However, increasing the contribution of viscous stresses from the surrounding 3D fluid(s) suppresses such an instability. We then tie this result to the hydrodynamic disturbances generated by each particle in the plane of the membrane and show that enhancing subphase viscous contributions generates extensional fields that orient neighboring particles in a manner that draws them apart. The balance of flux of particles aggregating versus separating then leads to a wave number selection in the mean-field model.

Biological membranes are self-assembled complex fluid interfaces that host proteins, molecular motors and other macromolecules essential for cellular function. These membranes have a distinct in-plane fluid response with a surface viscosity that has been well characterized. The resulting quasi-2D fluid dynamical problem describes the motion of embedded proteins or particles. However, the viscous response of biological membranes is often non-Newtonian: in particular, the surface shear viscosity of phospholipids that comprise the membrane depends strongly on the surface pressure. We use the Lorentz reciprocal theorem to extract the effective long-ranged hydrodynamic interaction among membrane inclusions that arises due to such non-trivial rheology. We show that the corrective force that emerges ties back to the interplay between membrane flow and non-constant viscosity, which suggests a mechanism for biologically favorable protein aggregation within membranes. We quantify and describe the mechanism for such a large-scale concentration instability using a mean-field model. Finally, we employ numerical simulations to demonstrate the formation of hexatic crystals due to the effective hydrodynamic interactions within the membrane.

Buckling plays a critical role in the transport and dynamics of elastic microfilaments in Stokesian fluids. However, previous work has only considered filaments with homogeneous structural properties. Filament backbone stiffness can be non-uniform in many biological systems like microtubules, where the association and disassociation of proteins can lead to spatial and temporal changes into structure. The consequences of such non-uniformities in the configurational stability and transport of these fibers are yet unknown. Here, we use slender-body theory and Euler-Bernoulli elasticity coupled with various non-uniform bending rigidity profiles to quantify this buckling instability using linear stability analysis and Brownian simulations. In shear flows, we observe more pronounced buckling in areas of reduced rigidity in our simulations. These areas of marked deformations give rise to differences in the particle extra stress, indicating a nontrivial rheological response due to the presence of these filaments. The fundamental mode shapes arising from each rigidity profile are consistent with the predictions from our linear stability analysis. Collectively, these results suggest that non-uniform bending rigidity can drastically alter fluid-structure interactions in physiologically relevant settings, providing a foundation to elucidate the complex interplay between hydrodynamics and the structural properties of biopolymers.

The surface shear viscosity of an insoluble surfactant monolayer often depends strongly on its surface pressure. Here, we show that a particle moving within a bounded monolayer breaks the kinematic reversibility of low-Reynolds-number flows. The Lorentz reciprocal theorem allows such irreversibilities to be computed without solving the full nonlinear equations, giving the leading-order contribution of surface-pressure-dependent surface viscosity. In particular, we show that a disk translating or rotating near an interfacial boundary experiences a force in the direction perpendicular to that boundary. In unbounded monolayers, coupled modes of motion can also lead to non-intuitive trajectories, which we illustrate using an interfacial analog of the Magnus effect. This perturbative approach can be extended to more complex geometries, and to 2D suspensions more generally.

In this fluid dynamics video we explore an array of different possible dynamics for a flexible filament sedimenting in a viscous fluid. The time-dependent shapes and trajectories of the filament are determined analytically and numerically by balancing viscous, elastic and gravitational forces in a slender-body theory for zero-Reynolds number flows. The dynamics are characterized by a single dimensionless elasto-gravitation number. The video shows the process of filament relaxation and reorientation, the formation of particle clouds, and finally a buckling instability., Comment: Fluid dynamics video entry for the Gallery of Fluid Motion at the 66th annual meeting of the American Physical Society - Division of Fluid Dynamics

The dynamics of a flexible filament sedimenting in a viscous fluid are explored analytically and numerically. Compared to the well-studied case of sedimenting rigid rods, the introduction of filament compliance is shown to cause a significant alteration in the long-time sedimentation orientation and filament geometry. A model is developed by balancing viscous, elastic, and gravitational forces in a slender-body theory for zero-Reynolds-number flows, and the filament dynamics are characterized by a dimensionless elasto-gravitation number. Filaments of both non-uniform and uniform cross-sectional thickness are considered. In the weakly flexible regime, a multiple-scale asymptotic expansion is used to obtain expressions for filament translations, rotations, and shapes. These are shown to match excellently with full numerical simulations. Furthermore, we show that trajectories of sedimenting flexible filaments, unlike their rigid counterparts, are restricted to a cloud whose envelope is determined by the elasto-gravitation number. In the highly flexible regime we show that a filament sedimenting along its long axis is susceptible to a buckling instability. A linear stability analysis provides a dispersion relation, illustrating clearly the competing effects of the compressive stress and the restoring elastic force in the buckling process. The instability travels as a wave along the filament opposite the direction of gravity as it grows and the predicted growth rates are shown to compare favorably with numerical simulations. The linear eigenmodes of the governing equation are also studied, which agree well with the finite-amplitude buckled shapes arising in simulations.