Stability of a biomembrane tube covered with proteins.

Membrane tubes are essential structural features in cells that facilitate biomaterial transport and inter- and intracellular signaling. The shape of these tubes can be regulated by the proteins that surround and adhere to them. We study the stability of a biomembrane tube coated with proteins by combining linear stability analysis, out-of-equilibrium hydrodynamic calculations, and numerical solutions of a Helfrich-like membrane model. Our analysis demonstrates that both long- and short-wavelength perturbations can destabilize the tubes. Numerical simulations confirm the derived linear stability criteria and yield the nonlinearly perturbed vesicle shapes. Our study highlights the interplay between membrane shape and protein density, where the shape instability concurs with a redistribution of proteins into a banded pattern.

Active flows and deformable surfaces in development.

We review progress in active hydrodynamic descriptions of flowing media on curved and deformable manifolds: the state-of-the-art in continuum descriptions of single-layers of epithelial and/or other tissues during development. First, after a brief overview of activity, flows and hydrodynamic descriptions, we highlight the generic challenge of identifying the dependence on dynamical variables of so-called active kinetic coefficients- active counterparts to dissipative Onsager coefficients. We go on to describe some of the subtleties concerning how curvature and active flows interact, and the issues that arise when surfaces are deformable. We finish with a broad discussion around the utility of such theories in developmental biology. This includes limitations to analytical techniques, challenges associated with numerical integration, fitting-to-data and inference, and potential tools for the future, such as discrete differential geometry.

Measuring Gaussian Rigidity Using Curved Substrates.

The Gaussian (saddle splay) rigidity of fluid membranes controls their equilibrium topology but is notoriously difficult to measure. In lipid mixtures, typical of living cells, linear interfaces separate liquid ordered (LO) from liquid disordered (LD) bilayer phases at subcritical temperatures. Here, we consider such membranes supported by curved substrates that thereby control the membrane curvatures. We show how spectral analysis of the fluctuations of the LO-LD interface provides a novel way of measuring the difference in Gaussian rigidity between the two phases. We provide a number of conditions for such interface fluctuations to be both experimentally measurable and sufficiently sensitive to the value of the Gaussian rigidity, while remaining in the perturbative regime of our analysis.

Dynamics of passive and active membrane tubes.

Utilising Onsager's variational formulation, we derive dynamical equations for the relaxation of a fluid membrane tube in the limit of small deformation, allowing for a contrast of solvent viscosity across the membrane and variations in surface tension due to membrane incompressibility. We compute the relaxation rates, recovering known results in the case of purely axis-symmetric perturbations and making new predictions for higher order (azimuthal) m-modes. We analyse the long and short wavelength limits of these modes by making use of various asymptotic arguments. We incorporate stochastic terms to our dynamical equations suitable to describe both passive thermal forces and non-equilibrium active forces. We derive expressions for the fluctuation amplitudes, an effective temperature associated with active fluctuations, and the power spectral density for both the thermal and active fluctuations. We discuss an experimental assay that might enable measurement of these fluctuations to infer the properties of the active noise. Finally we discuss our results in the context of active membranes more generally and give an overview of some open questions in the field.

Shear-Driven Instabilities of Membrane Tubes and Dynamin-Induced Scission.

Motivated by the mechanics of dynamin-mediated membrane tube fission, we analyze the stability of fluid membrane tubes subjected to shear flow in azimuthal direction. We find a novel helical instability driven by the membrane shear flow which results in a nonequilibrium steady state for the tube fluctuations. This instability has its onset at shear rates that may be physiologically accessible under the action of dynamin and could also be probed using in vitro experiments on membrane nanotubes, e.g., using magnetic tweezers. We discuss how such an instability may play a role in the mechanism for dynamin-mediated membrane tube fission.

Hydro-osmotic Instabilities in Active Membrane Tubes.

We study a membrane tube with unidirectional ion pumps driving an osmotic pressure difference. A pressure-driven peristaltic instability is identified, qualitatively distinct from similar tension-driven Rayleigh-type instabilities on membrane tubes. We discuss how this instability could be related to the function and biogenesis of membrane bound organelles, in particular, the contractile vacuole complex. The unusually long natural wavelength of this instability is in agreement with that observed in cells.