Active matter systems such as eukaryotic cells and bacteria continuously transform chemical energy to motion. Hence living systems exert active stresses on the complex environments in which they reside. One recurring aspect of this complexity is the viscoelasticity of the medium surrounding living systems: bacteria secrete their own viscoelastic extracellular matrix, and cells constantly deform, proliferate, and self-propel within viscoelastic networks of collagen. It is therefore imperative to understand how active matter modifies, and gets modified by, viscoelastic fluids. Here, we present a two-phase model of active nematic matter that dynamically interacts with a passive viscoelastic polymeric phase and perform numerical simulations in two dimensions to illustrate its applicability. Motivated by recent experiments we first study the suppression of cell division by a viscoelastic medium surrounding the cell. We further show that the self-propulsion of a model keratocyte cell is modified by the polymer relaxation of the surrounding viscoelastic fluid in a non-uniform manner and find that increasing polymer viscosity effectively suppresses the cell motility. Lastly, we explore the hampering impact of the viscoelastic medium on the generic hydrodynamic instabilities of active nematics by simulating the dynamics of an active stripe within a polymeric fluid. The model presented here can provide a framework for investigating more complex dynamics such as the interaction of multicellular growing systems with viscoelastic environments.
Swimming microorganisms often self propel in fluids with complex rheology. While past theoretical work indicates that fluid viscoelasticity should hinder their locomotion, recent experiments on waving swimmers suggest a possible non-Newtonian enhancement of locomotion. We suggest a physical mechanism, based on fluid-structure interaction, leading to swimming in a viscoelastic fluid at a higher speed than in a Newtonian one. Using Taylor's two-dimensional swimming sheet model, we solve for the shape of an active swimmer as a balance between the external fluid stresses, the internal driving mome…
We derive a general formula for the inertialess dynamics of active particles in linear viscoelastic fluids by means of a modified reciprocal theorem. We then demonstrate that force-free active particles in Maxwell-like linear viscoelastic fluids with no retardation have exactly the same dynamics as in Newtonian fluids. In contrast, active particles in Jeffreys-like fluids with retardation can display markedly different dynamics, including net motion under reciprocal actuation thereby breaking the scallop theorem.
Complex interactions between cellular systems and their surrounding extracellular matrices are emerging as important mechanical regulators of cell functions such as proliferation, motility, and cell death, and such cellular systems are often characterized by pulsating acto-myosin activities. Here, using an active gel model, we numerically explore the spontaneous flow generation by activity pulses in the presence of a viscoelastic medium. The results show that cross-talk between the activity-induced deformations of the viscoelastic surroundings with the time-dependent response of the active med…
Active matter consists of units that generate mechanical work by consuming energy. Examples include living systems, such as assemblies of bacteria and biological tissues, biopolymers driven by molecular motors, and suspensions of synthetic self-propelled particles. A central question in the field is to understand and control the self-organization of active assemblies in space and time. Most active systems exhibit either spatial order mediated by interactions that coordinate the spatial structure and the motion of active agents or the temporal synchronization of individual oscillatory dynamics.…
We show that a viscoelastic thin sheet driven out of equilibrium by active structural remodelling develops a rich variety of shapes as a result of a competition between viscous relaxation and activity. In the regime where active processes are faster than viscoelastic relaxation, wrinkles that are formed due to remodelling are unable to relax to a configuration that minimises the elastic energy and the sheet is inherently out of equilibrium. We argue that this non-equilibrium regime is of particular interest in biology as it allows the system to access morphologies that are unavailable if restr…
Models of active nematics in biological systems normally require complexity arising from the hydrodynamics involved at the microscopic level as well as the viscoelastic nature of the system. Here we show that a minimal, space-independent, model based on the temporal alignment of active and polymeric particles provides an avenue to predict and study their coupled dynamics within the framework of dynamical systems. In particular, we examine, using analytical and numerical methods, how such a simple model can display self-sustained oscillations in an activity-driven viscoelastic shear flow.
Constitutive equations for a one-dimensional, active, polar, viscoelastic liquid are derived by treating the strain field as a slow hydrodynamic variable. Taking into account the couplings between strain and polarity allowed by symmetry, the hydrodynamics of an active, polar, viscoelastic body include an evolution equation for the polarity field that generalizes the damped Kuramoto-Sivashinsky equation. Beyond thresholds of the active coupling coefficients between the polarity and the stress or the strain rate, bifurcations of the homogeneous state lead first to stationary waves, then to propa…
Based on a Hamiltonian that incorporates the elastic coupling between a tracer and active particles, we derive a generalized Langevin model for the non-equilibrium mechanical response of active viscoelastic biomatter. Our model accounts for the power-law viscoelastic response of the embedding polymeric network as well as for the non-equilibrium energy transfer between active and tracer particles. Our analytical expressions for the frequency-dependent response function and the positional autocorrelation function agree nicely with experimental data for red blood cells and actomyosin networks wit…
The mechanical response of active media ranging from biological gels to living tissues is governed by a subtle interplay between viscosity and elasticity. In this Letter, we generalize the canonical Kelvin-Voigt and Maxwell models to active viscoelastic media that break both parity and time-reversal symmetries. The resulting continuum theories exhibit viscous and elastic tensors that are both antisymmetric, or odd, under exchange of pairs of indices. We analyze how these parity violating viscoelastic coefficients determine the relaxation mechanisms and wave-propagation properties of odd materi…
We have directly observed short-time stress propagation in viscoelastic fluids using two optically trapped particles and a fast interferometric particle-tracking technique. We have done this both by recording correlations in the thermal motion of the particles and by measuring the response of one particle to the actively oscillated second particle. Both methods detect the vortex-like flow patterns associated with stress propagation in fluids. This inertial vortex flow propagates diffusively for simple liquids, while for viscoelastic solutions the pattern spreads super-diffusively, dependent on…
We consider a continuum model of active viscoelastic matter, whereby an active nematic liquid-crystal is coupled to a minimal model of polymer dynamics with a viscoelastic relaxation time $τ_C$. To explore the resulting interplay between active and polymeric dynamics, we first generalise a linear stability analysis (from earlier studies without polymer) to derive criteria for the onset of spontaneous heterogeneous flows (strain rate) and/or deformations (strain). We find two modes of instability. The first is a viscous mode, associated with strain rate perturbations. It dominates for relativel…
A dilute suspension of active Brownian particles in a dense compressible viscoelastic fluid, forms a natural setting to study the emergence of nonreciprocity during a dynamical phase transition. At these densities, the transport of active particles is strongly influenced by the passive medium and shows a dynamical jamming transition as a function of activity and medium density. In the process, the compressible medium is actively churned up -for low activity, the active particle gets self-trapped in a spherical cavity of its own making, while for large activity, the active particle ploughs thro…
In this work, we investigate the dynamics of the number density fluctuations of a dilute suspension of active particles in a linear viscoelastic fluid. We propose a model for the frequency-dependent diffusion coefficient of the active particles, which captures the effect of rotational diffusion on the persistence of their self-propelled motion and the viscoelasticity of the medium. Using fluctuating hydrodynamics, the linearized equations for the active suspension are derived, from which we calculate its dynamic structure factor and the corresponding intermediate scattering function. For a Max…
By now active Brownian motion is a well-established model to describe the motion of mesoscopic self-propelled particles in a Newtonian fluid. On the basis of the generalized Langevin equation, we present an analytic framework for active Brownian motion with memory delay assuming time-dependent friction kernels for both translational and orientational degrees of freedom to account for the time-delayed response of a viscoelastic medium. Analytical results are obtained for the orientational correlation function, mean displacement and mean-square displacement which we evaluate in particular for a …
Active chiral viscoelastic materials exhibit elastic responses perpendicular to the applied stresses, referred to as odd elasticity. We use a covariant formulation of viscoelasticity combined with an entropy production analysis to show that odd elasticity is not only present in active systems but also in broad classes of passive chiral viscoelastic fluids. In addition, we demonstrate that linear viscoelastic chiral solids do require activity in order to manifest odd elastic responses. In order to model the phenomenon of passive odd viscoelasticity we propose a chiral extension of Jeffreys mode…
We experimentally study the dynamics of active particles (APs) in a viscoelastic fluid under various geometrical constraints such as flat walls, spherical obstacles and cylindrical cavities. We observe that the main effect of the confined viscoelastic fluid is to induce an effective repulsion on the APs when moving close to a rigid surface, which depends on the incident angle, the surface curvature and the particle activity. Additionally, the geometrical confinement imposes an asymmetry to their movement, which leads to strong hydrodynamic torques, thus resulting in detention times on the wall…
With the use of the "two-fluid model", we discuss anomalous diffusion induced by active force dipoles in viscoelastic media. Active force dipoles, such as proteins and bacteria, generate non-thermal fluctuating flows that lead to a substantial increment of the diffusion. Using the partial Green's function of the two-fluid model, we first obtain passive (thermal) two-point correlation functions such as the displacement cross-correlation function between the two point particles separated by a finite distance. We then calculate active (non-thermal) one-point and two-point correlation functions du…
Run-and-tumble (RNT) motion is a prominent locomotion strategy employed by many living microorganisms. It is characterized by straight swimming intervals (runs), which are interrupted by sudden reorientation events (tumbles). In contrast, directional changes of synthetic microswimmers (active particles, APs) are caused by rotational diffusion, which is superimposed with their translational motion and thus leads to rather continuous and slow particle reorientations. Here we demonstrate that active particles can also perform a swimming motion where translational and orientational changes are dis…
A paradigm for internally driven matter is the active nematic liquid crystal, whereby the equations of a conventional nematic are supplemented by a minimal active stress that violates time reversal symmetry. In practice, active fluids may have not only liquid crystalline but also viscoelastic polymer degrees of freedom. Here we explore the resulting interplay by coupling an active nematic to a minimal model of polymer rheology. We find that adding polymer can greatly increase the complexity of spontaneous flow, but can also have calming effects, thereby increasing the net throughput of spontan…
Beyond the Markovian limit: Exact solutions for active motion in a power-law viscoelastic bath
Wenzhou Institute of the University of Chinese Academy of Sciences（中国科学院大学温州研究院）；School of Physical Sciences, University of Chinese Academy of Sciences（中国科学院大学物理科学学院）；Departament de Física de la Matèria Condensada, Universitat de Barcelona（巴塞罗那大学凝聚态物理学系）；Chinese Academy of Sciences Key Laboratory of Soft Matter Physics, Institute of Physics, Chinese Academy of Sciences（中国科学院物理研究所软物质物理重点实验室）；UBICS University of Barcelona Institute of Complex Systems（巴塞罗那大学复杂系统研究所）
Viscoelastic liquids are characterized by a finite static viscosity and a zero yield stress, whereas soft solids have an infinite viscosity and a non-zero yield stress. The rheological nature of viscoelastic materials has long been a challenge, and it is still a matter of debate. Here, we provide for the first time the constitutive equations of linear viscoelasticity for magnetic wires in yield stress materials, together with experimental measurements using Magnetic Rotational Spectroscopy (MRS). With MRS, the wires are submitted to a rotational magnetic field as a function of frequency and th…
We use active microrheology to determine the frequency dependent moduli of a linear viscoelastic fluid in terms of the polymer time constant ($λ$), and the polymer ($μ_p$) and solvent viscosity ($μ_s$), respectively. We measure these parameters from the response function of an optically trapped Brownian probe in the fluid under an external perturbation, and at different dilutions of the viscoelastic component in the fluid. This is an improvement over bulk microrheology measurements in viscoelastic Stokes-Oldroyd B fluids which determine the complex elastic modulus $G(ω)$ of the fluid, but do n…
We consider the dynamics of a charged inertial active Ornstein-Uhlenbeck particle in a viscoelastic suspension under the action of an uniform magnetic field. With the help of both numerical simulation and analytical framework, we exactly investigate the viscoelastic response of the particle to the magnetic field by means of particle trajectories, mean displacement, and mean square displacement (MSD) calculations. The simulated particle trajectories and MSD calculations reveal that the steady state response of a confined harmonic particle to the magnetic field show interesting features due to t…
Active wetting/de-wetting of focal adhesions on viscoelastic substrates
University of Belgrade（贝尔格莱德大学）；Victor Chang Cardiac Research Institute（维克多·张心脏研究所）；University of New South Wales（新南威尔士大学）；University of Glasgow（格拉斯哥大学）；Lancaster University（兰开斯特大学）
We investigate a model for a Stirling-like engine consisting of a passive Brownian particle confined by a harmonic potential and interacting with a suspension of active Brownian particles that self-propel in a viscous solvent, which cyclically operates under isothermal conditions by means of temporal variations of the trap stiffness and the self-propulsion speed of the active particles. We derive an effective stochastic equation of motion of the trapped Brownian particle, which includes a friction memory kernel as well as thermal and active fluctuating forces due to its coupling with the activ…
Optical tracking in vivo experiments reveal that diffusion of particles in biological cells is strongly enhanced in the presence of ATP and the experimental data for animal cells could previously be reproduced within a phenomenological model of a gel with myosin motors acting within it [EPL 110, 48005 (2015)]. Here, the two-fluid model of a gel is considered where active macromolecules, described as force dipoles, cyclically operate both in the elastic and the fluid components. Through coarse-graining, effective equations of motions for tracer particles displaying local deformations and local …
Emergent odd viscoelasticity in chiral soft glassy materials
University of Göttingen（哥廷根大学）；Ecole Polytechnique de l’Université de Nantes, LTeN, U6607, Nantes Université（南特大学理工学院，LTeN，U6607）；King’s College London（伦敦国王学院）
Many wormlike micellar systems exhibit appreciable shear thinning due to shear induced alignment. As the micelles get aligned introducing directionality in the system, the viscoelastic properties are no longer expected to be isotropic. An optical tweezers based active microrheology technique enables us to probe the out-of-equilibrium rheological properties of a wormlike micellar system simultaneously along two orthogonal directions - parallel to the applied shear, as well as perpendicular to it. While the displacements of a trapped bead - in response to active drag force carry signature of con…
Inertial cavitation in soft matter is an important phenomenon featured in a wide array of biological and engineering processes. Recent advances in experimental, theoretical, and numerical techniques have provided access into a world full of nonlinear physics, yet most of our quantitative understanding to date has been centered on a spherically symmetric description of the cavitation process. However, cavitation bubble growth and collapse rarely occur in a perfectly symmetrical fashion, particularly in soft materials. Predicting the onset of dynamically arising, non-spherical instabilities has …
The onset of rigidity in interacting liquids, as they undergo a transition to a disordered solid, is associated with a rearrangement of the low-frequency vibrational spectrum. In this letter, we derive scaling forms for the singular dynamical response of disordered viscoelastic networks near both jamming and rigidity percolation. Using effective-medium theory, we extract critical exponents, invariant scaling combinations and analytical formulas for universal scaling functions near these transitions. Our scaling forms describe the behavior in space and time near the various onsets of rigidity, …
Many swimming microorganisms, such as bacteria and sperm, use flexible flagella to move through viscoelastic media in their natural environments. In this paper we address the effects a viscoelastic fluid has on the motion and beating patterns of elastic filaments. We treat both a passive filament which is actuated at one end, and an active filament with bending forces arising from internal motors distributed along its length. We describe how viscoelasticity modifies the hydrodynamic forces exerted on the filaments, and how these modified forces affect the beating patterns. We show how high vis…
Many problems in mechanobiology urgently require characterisation of the micromechanical properties of the fibrous proteins of cells and tissues. Brillouin light scattering has been proposed as a new optical elastography technique to meet this need, but the information contained in the Brillouin spectrum is still a matter of debate. Here we investigate this question using gelatin as a model system in which the macroscopic physical properties can be manipulated to mimic all the relevant biological states of matter, ranging from the liquid to the gel and the glassy phase. We demonstrate that Bri…
Constitutive equations are derived for the viscoelastic response of rubbery polymers at finite strains. A polymer is thought of as a network of long chains connected to temporary junctions. At a random time, a chain detaches from a junction, which is treated at transition from its active state to the dangling state. A dangling chain randomly captures a new junction in the vicinity of its free end and returns to its active state. Breakage and reformation of long chains are modeled as thermo-mechanically activated processes. Stress-strain relations for a rubbery polymer are developed using the l…
Disordered viscoelastic materials are ubiquitous and exhibit fascinating invariant scaling properties. In a companion article, we have presented comprehensive new results for the critical behavior of the dynamic susceptibility of disordered elastic systems near the onset of rigidity. Here we provide additional details of the derivation of the singular scaling forms of the longitudinal response near both jamming and rigidity percolation. We then discuss global aspects associated with these forms, and make scaling collapse plots for both undamped and overdamped dynamics in both the rigid and flo…
Characterizing the softness of deformable materials having partial elastic and partial viscous behaviour via soft lubrication experiments has emerged as a versatile and robust methodology in recent times. However, a straightforward extension of the classical elastohydrodynamic lubrication theory that is commonly employed for characterizing elastic materials turns out to be rather inadequate in explaining the response of such viscoelastic materials subjected to dynamic loading conditions, despite adhering to a mathematically acceptable framework via the complex Young's modulus as a material pro…
Many practically relevant materials combine properties of viscous fluids and elastic solids to viscoelastic behavior. Our focus is on the induced dynamic behavior of damped finite-sized particulate inclusions in such substances. We explicitly describe history-dependent interactions that emerge between the embedded particles. These interactions are mediated by the viscoelastic surroundings. They result from the flows and distortions of the viscoelastic medium when induced by the rigid inclusions. Both, viscoelastic environments of terminal fluid-like flow or of completely reversible damped elas…
