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Sami C. Al-Izzi

Publications and source records attributed to Sami C. Al-Izzi.

13 recordsLinked to original sources

Tuning nonlinear waves in nonreciprocal active filaments

The instabilities of slender structures power biological locomotion across scales, and offer a compelling method to actuate soft robots. Nonreciprocal elastic solids have been found to amplify flexural waves in one direction only, but design principles to tune and stabilize these waves are missing. Here we develop a geometrically exact theory of nonreciprocal filaments and provide simulations that capture their post-instability nonlinear dynamics. We find that nonreciprocity, when coupled to inertia or pre-stress, amplifies and advects curvature variations. The resulting one-way patterns of shape morphing can then be selected via dissipative interactions with the environment. Our work offers a continuum-based strategy for how internal stresses can drive active unidirectional waves without need for additional degrees of freedom.

cond-mat.soft

Nonreciprocal buckling makes active filaments polyfunctional

Active filaments are a workhorse for propulsion and actuation across biology, soft robotics and mechanical metamaterials. However, artificial active rods suffer from limited robustness and adaptivity because they rely on external control, or are tethered to a substrate. Here we bypass these constraints by demonstrating that non-reciprocal interactions lead to large-scale unidirectional dynamics in free-standing slender structures. By coupling the bending modes of a buckled beam anti-symmetrically, we transform the multistable dynamics of elastic snap-through into persistent cycles of shape change. In contrast to the critical point underpinning beam buckling, this transition to self-snapping is mediated by a critical exceptional point, at which bending modes simultaneously become unstable and degenerate. Upon environmental perturbation, our active filaments exploit self-snapping for a range of functionality including crawling, digging and walking. Our work advances critical exceptional physics as a guiding principle for programming instabilities into functional active materials.

cond-mat.soft

Capillarity Reveals the Role of Capsid Geometry in HIV Nuclear Translocation

The protective capsid encasing the genetic material of Human Immunodeficiency Virus (HIV) has been shown to traverse the nuclear pore complex (NPC) intact, despite exceeding the passive diffusion threshold by over three orders of magnitude. This remarkable feat is attributed to the properties of the capsid surface, which confer solubility within the NPC's phase-separated, condensate-like barrier. In this context, we apply the classical framework of wetting and capillarity -- integrating analytical methods with sharp- and diffuse-interface numerical simulations -- to elucidate the physical underpinnings of HIV nuclear entry. Our analysis captures several key phenomena: the reorientation of incoming capsids due to torques arising from asymmetric capillary forces; the role of confinement in limiting capsid penetration depths; the classification of translocation mechanics according to changes in topology and interfacial area; and the influence of (spontaneous) rotational symmetry-breaking on energetics. These effects are all shown to depend critically on capsid geometry, arguing for a physical basis for HIV's characteristic capsid shape.

physics.bio-ph

More ATP Does Not Equal More Contractility: Power And Remodelling In Reconstituted Actomyosin

The cytoskeletal component actomyosin is a canonical example of active matter since the powerstroke cycle locally converts chemical energy in the form of adenoside triphosphate (ATP) into mechanical work for remodelling. Observing myosin II minifilaments as they remodel actin {\it in vitro}, we now report that: at high concentrations of ATP, myosin minifilaments form metastable swirling patterns that are characterised by recurrent vortex and spiral-like motifs, whereas; at low concentrations of ATP, such structures give way to aster-like patterns. To explain this, we construct the (quasi-)steady states of a polar active hydrodynamic theory of actomyosin whose ATP-scaling is obtained from a microscopic, stochastic description for the ATP-dependent binding of the heads of single myosin II minifilaments. The latter codifies the heuristic that, since the powerstroke cycle involves the unbinding of myosin II heads from actin, increases in the concentration of ATP reduce the likelihood that a given myosin II minifilament has more than one head bound simultaneously, reducing its ability to generate contractile forces and increasing the relative likelihood of processive motion. This reproduces several qualitative and some quantitative aspects of experiments, providing evidence for the central phenomenon of the theory: an ATP-dependent active contractile instability. ATP therefore controls not only the rate at which work is done -- \textit{i.e.,} the power -- but also the mode by which this occurs.

physics.bio-ph

Membrane tubes with active pumping: water transport, vacuole formation and osmoregulation

The need for organisms to regulate their volume and osmolarity when surrounded by freshwater is a basic physical challenge for many bacteria, protists and algae. Taking inspiration from the contractile vacuole complex found in many protists, we discuss how simple models of active membrane tubes can give insights into the fluid and active ionic transport properties of such systems. We show that a simple membrane tube with unidirectional ion pumps, and passive ion and water channels, forms a large vacuole due to osmotically-driven water flow and that this can be used to actively pump water out of the cell interior. We discuss the use of this system as a possible minimal method for osmoregulation.

physics.bio-ph

Advecting Scaffolds: Controlling The Remodelling Of Actomyosin With Anillin

We propose and analyse an active hydrodynamic theory that characterises the effects of the scaffold protein anillin. Anillin is found at major sites of cortical activity, such as adherens junctions and the cytokinetic furrow, where the canonical regulator of actomyosin remodelling is the small GTPase, RhoA. RhoA acts via intermediary 'effectors' to increase both the rates of activation of myosin motors and the polymerisation of actin filaments. Anillin has been shown to scaffold this action of RhoA - improving critical rates in the signalling pathway without altering the essential biochemistry - but its contribution to the wider spatio-temporal organisation of the cortical cytoskeleton remains poorly understood. Here, we combine analytics and numerics to show how anillin can non-trivially regulate the cytoskeleton at hydrodynamic scales. At short times, anillin can amplify or dampen existing contractile instabilities, as well as alter the parameter ranges over which they occur. At long times, it can change both the size and speed of steady-state travelling pulses. The primary mechanism that underpins these behaviours is established to be the advection of anillin by myosin II motors, with the specifics relying on the values of two coupling parameters. These codify anillin's effect on local signalling kinetics and can be traced back to its interaction with the acidic phospholipid phosphatidylinositol 4,5-bisphosphate (PIP2), thereby establishing a putative connection between actomyosin remodelling and membrane composition.

q-bio.SC

A Twist On Active Membranes: Odd Mechanics, Spontaneous Flows and Shape Instabilities

Living systems are chiral on multiple scales, from constituent biopolymers to large scale morphology, and their active mechanics is both driven by chiral components and serves to generate chiral morphologies. We describe the mechanics of active fluid membranes in coordinate-free form, with focus on chiral contributions to the stress. These generate geometric `odd elastic' forces in response to mean curvature gradients but directed perpendicularly. As a result, they induce tangential membrane flows that circulate around maxima and minima of membrane curvature. When the normal viscous force amplifies perturbations the membrane shape can become linearly unstable giving rise to shape instabilities controlled by an active Scriven-Love number. We describe examples for spheroids, membranes tubes and helicoids, discussing the relevance and predictions such examples make for a variety of biological systems from the sub-cellular to tissue level.

cond-mat.soft

Morphodynamics of Active Nematic Fluid Surfaces

Morphodynamic equations governing the behaviour of active nematic fluids on deformable curved surfaces are constructed in the large deformation limit. Emphasis is placed on the formulation of objective rates that account for normal deformations whilst ensuring that tangential flows are Eulerian, and the use of the surface derivative (rather than the covariant derivative) in the nematic free energy, which elastically couples local order to out-of-plane bending of the surface. Focusing on surface geometry and its dynamical interplay with the hydrodynamics, several illustrative instabilities are then characterised. These include cases where the role of the Scriven-Love number and its nematic analogue are non-negligible, and where the active nematic forcing can be characterised by an analogue of the Föppl-von-Kármán number. For the former, flows and changes to the nematic texture are coupled to surface geometry by viscous dissipation. This is shown to result in non-trivial relaxation dynamics for a nematic tube. For the latter, the nematic active forcing couples to the surface bending terms of the nematic free energy, resulting in extensile (active ruffling) and contractile (active pearling) instabilities in the tube shape, as well as active bend instabilities in the nematic texture. In comparison to the flat case, such bend instabilities now have a threshold set by the extrinsic curvature of the tube. Finally, we examine a topological defect located on an almost flat surface and show that there exists a steady state where a combination of defect elasticity, activity and non-negligible spin-connection drive a shape change in the surface.

cond-mat.soft

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.

physics.bio-ph

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 supports 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, whilst remaining in the perturbative regime of our analysis.

cond-mat.soft

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.

cond-mat.soft

Shear-driven instabilities of membrane tubes and dynamin-induced scission

Motivated by the mechanics of dynamin-mediated membrane tube fission we analyse 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 non-equilibrium 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.

physics.bio-ph

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.

physics.bio-ph