arXiv subjects

Raphael Voituriez

Publications and source records attributed to Raphael Voituriez.

At least 19 recordsLinked to original sources

arXiv2026

Density and shape govern the dynamical self-organization of active matter on a droplet

Morphogenesis emerges from dynamic feedback among geometry, mechanics, and chemistry; however, disentangling these contributions in living systems remains challenging. Here, we focus on the interplay between geometry and mechanics by developing a minimal in vitro model in which purified microtubules and kinesin motor clusters self-organize into a two-dimensional active nematic cortex at the surface of spherical water-in-oil droplets. The spherical geometry enforces a total topological charge of +2, here realized by four +1/2 defects whose trajectories reveal robust, self-sustained oscillations. Using full-surface reconstructions, we show that the collective dynamics of the defects lead to a periodic switching between planar and tetrahedral arrangements through alternating coiling and hemisphere-crossing phases. By tuning microtubule density, the system spans a continuum from a classic defect-dominated active nematic to a regime resembling an extensile filament confined to a curved surface, where low density is associated with increased trajectory variability and direction reversals. Geometric perturbations introduced through controlled squeezing redistribute curvature and induce the nucleation of additional defects, thereby reorganizing the entire topological landscape while preserving total charge. Together, these results show that periodic morphogenetic-like cycles, defect topology, and material organization can arise solely from the interplay of activity, density, and curvature. This reconstituted system provides a versatile platform for elucidating the coupling between mechanics and geometry underlying shape formation in active biological matter.

cond-mat.soft↗

arXiv2025

Stochastic Forces Enhance Tracer Diffusion in Non-motile Active Matter

Stochasticity is a defining feature of the pairwise forces governing interactions in biological systems-from molecular motors to cell-cell adhesion-yet its consequences on large-scale dynamics remain poorly understood. Here, we show that reciprocal but randomly fluctuating interactions between particles create active suspensions which can enhance the diffusion of an external tracer particle, even in the absence of self-propulsion or non-reciprocity. Starting from a lattice model with pairwise dynamics that minimally break detailed balance, we derive a coarse-grained dynamical theory for spatio-temporal density fluctuations and reveal an elevated effective temperature at short wavelengths. We then compute the self-diffusion coefficient of a tracer particle weakly coupled to our active fluid, demonstrating that purely reciprocal stochastic interactions provide a distinct and generic route to enhanced diffusivity in dense non-equilibrium suspensions.

cond-mat.soft↗

arXiv2025

Adhesion differentials control the rheology of biomimetic emulsions

Animal morphogenesis involves complex tissue deformation processes, which require tight control over tissue rheology. Yet, it remains insufficiently understood how tissue rheology results from the interplay between cellular packing and cellular forces, such as cortical tension, cell pressure, and cell-cell adhesion. Here, we follow a biomimetic approach to study this interplay. We mimic adhesive cells with oil droplets whose adhesion strength and specificity can be flexibly tuned. Using microfluidics, we expose 2D emulsions to an oscillatory geometry imposing cyclic pure shear, and we develop a geometric method to quantify their rheology using only imaging data. We find that some of the emulsions made of two droplet types progressively change their yielding behavior across subsequent shear cycles. Combining this with vertex model simulations, we show that the observed shift in yielding behavior is due to a progressive compaction, which only occurs in emulsions with a high adhesion differential and only when exposed to oscillatory shear. Gradients of cell compaction have been observed during animal development. Our work demonstrates how such gradients can be used to control gradients of tissue rheological properties. Moreover, the progressive compaction suggests the emergence of a pumping mechanism, which potentially acts in many cellular materials, from foams to tissues.

cond-mat.soft↗

arXiv2024

Scale-free flocking and giant fluctuations in epithelial active solids

The collective motion of epithelial cells is a fundamental biological process which plays a significant role in embryogenesis, wound healing and tumor metastasis. While it has been broadly investigated for over a decade both in vivo and in vitro, large scale coherent flocking phases remain underexplored and have so far been mostly described as fluid. In this work, we report a mode of large-scale collective motion for different epithelial cell types in vitro with distinctive new features. By tracking individual cells, we show that cells move over long time scales coherently not as a fluid, but as a polar elastic solid with negligible cell rearrangements. Our analysis reveals that this solid flocking phase exhibits signatures of long-range polar order, unprecedented in cellular systems, such as scale-free correlations, anomalously large density fluctuations, and shear waves. Based on a general theory of active polar solids, we argue that these features result from massless Goldstone modes, which, in contrast to polar fluids where they are generic, require the decoupling of global rotations of the polarity and in-plane elastic deformations in polar solids. We theoretically show and consistently observe in experiments that the fluctuations of elastic deformations diverge for large system size in such polar active solid phases, leading eventually to rupture and thus potentially loss of tissue integrity at large scales.

cond-mat.soft↗

arXiv2024

Ordering, spontaneous flows and aging in active fluids depositing tracks

Growing experimental evidence shows that cell monolayers can induce long-lived perturbations to their environment, akin to footprints, which in turn influence the global dynamics of the system. Inspired by these observations, we propose a comprehensive theoretical framework to describe systems where an active field dynamically interacts with a non-advected footprint field, deposited by the active field. We derive the corresponding general hydrodynamics for both polar and nematic fields. Our findings reveal that the dynamic coupling to a footprint field induces remarkable effects absent in classical active hydrodynamics, such as symmetry-dependent modifications to the isotropic-ordered transition, which may manifest as either second-order or first-order, alterations in spontaneous flow transitions, potentially resulting in oscillating flows and rotating fields, and initial condition-dependent aging dynamics characterized by long-lived transient states. Our results suggest that footprint deposition could be a key mechanism determining the dynamical phases of cellular systems, or more generally active systems inducing long-lived perturbations to their environment.

cond-mat.soft↗

arXiv2023

Aging Dynamics of $d-$dimensional Locally Activated Random Walks

Locally activated random walks are defined as random processes, whose dynamical parameters are modified upon visits to given activation sites. Such dynamics naturally emerge in living systems as varied as immune and cancer cells interacting with spatial heterogeneities in tissues, or at larger scales animals encountering local resources. At the theoretical level, these random walks provide an explicit construction of strongly non Markovian, and aging dynamics. We propose a general analytical framework to determine various statistical properties characterizing the position and dynamical parameters of the random walker on $d$-dimensional lattices. Our analysis applies in particular to both passive (diffusive) and active (run and tumble) dynamics, and quantifies the aging dynamics and potential trapping of the random walker; it finally identifies clear signatures of activated dynamics for potential use in experimental data.

cond-mat.stat-mech↗

arXiv2022

Crosslinking and depletion determine spatial instabilities in cytoskeletal active matter

Active gels made of cytoskeletal proteins are valuable materials with attractive non-equilibrium properties such as spatial self-organization and self-propulsion. At least four typical routes to spatial patterning have been reported to date in different types of cytoskeletal active gels: bending and buckling instabilities in extensile systems, and global and local contraction instabilities in contractile gels. Here we report the observation of these four instabilities in a single type of active gel and we show that they are controlled by two parameters: the concentrations of ATP and depletion agent. We demonstrate that as the ATP concentration decreases, the concentration of passive motors increases until the gel undergoes a gelation transition. At this point, buckling is selected against bending, while global contraction is favored over local ones. Our observations are coherent with a hydrodynamic model of a viscoelastic active gel where the filaments are crosslinked with a characteristic time that diverges as the ATP concentration decreases. Our work thus provides a unified view of spatial instabilities in cytoskeletal active matter.

cond-mat.soft↗

arXiv2021

Self-interacting random walks : aging, exploration and first-passage times

Self-interacting random walks are endowed with long range memory effects that emerge from the interaction of the random walker at time $t$ with the territory that it has visited at earlier times $t'<t$. This class of non Markovian random walks has applications in a broad range of examples, ranging from insects to living cells, where a random walker modifies locally its environment -- leaving behind footprints along its path, and in turn responds to its own footprints. Because of their inherent non Markovian nature, the exploration properties of self-interacting random walks have remained elusive. Here we show that long range memory effects can have deep consequences on the dynamics of generic self-interacting random walks ; they can induce aging and non trivial persistence and transience exponents, which we determine quantitatively, in both infinite and confined geometries. Based on this analysis, we quantify the search kinetics of self-interacting random walkers and show that the distribution of the first-passage time (FPT) to a target site in a confined domain takes universal scaling forms in the large domain size limit, which we characterize quantitatively. We argue that memory abilities induced by attractive self-interactions provide a decisive advantage for local space exploration, while repulsive self-interactions can significantly accelerate the global exploration of large domains.

cond-mat.stat-mech↗

arXiv2020

Chiral active hexatics: Giant number fluctuations, waves and destruction of order

Active materials, composed of internally driven particles, have properties that are qualitatively distinct from matter at thermal equilibrium. However, the most spectacular departures from equilibrium phase behaviour are thought to be confined to systems with polar or nematic asymmetry. In this paper, we show that such departures are also displayed in more symmetric phases such as hexatics if, in addition, the constituent particles have chiral asymmetry. We show that chiral active hexatics whose rotation rate does not depend on density have giant number fluctuations. If the rotation rate depends on density, the giant number fluctuations are suppressed due to a novel orientation-density sound mode with a linear dispersion which propagates even in the overdamped limit. However, we demonstrate that beyond a finite but large lengthscale, a chirality and activity-induced relevant nonlinearity invalidates the predictions of the linear theory and destroys the hexatic order. In addition, we show that activity modifies the interactions between defects in the active chiral hexatic phase, making them non-mutual. Finally, to demonstrate the generality of a chiral active hexatic phase we show that it results from the melting of chiral active crystals in finite systems.

cond-mat.soft↗

arXiv2020

One dimensional cell motility patterns

During migration cells exhibit a rich variety of seemingly random migration patterns, which makes unraveling the underlying mechanisms that control cell migration a daunting challenge. For efficient migration cells require a mechanism for polarization, so that traction forces are produced in the direction of motion, while adhesion is released to allow forward migration. To simplify the study of this process cells have been studied when placed along one-dimensional tracks, where single cells exhibit both smooth and stick-slip migration modes. The stick-slip motility mode is characterized by protrusive motion at the cell front, coupled with slow cell elongation, which is followed by rapid retractions of the cell back. In this study, we explore a minimal physical model that couples the force applied on the adhesion bonds to the length variations of the cell and the traction forces applied by the polarized actin retrograde flow. We show that the rich spectrum of cell migration patterns emerges from this model as different \emph{deterministic} dynamical phases. This result suggests a source for the large cell-to-cell variability (CCV) in cell migration patterns observed in single cells over time and within cell populations. The large heterogeneity can arise from small fluctuations in the cellular components that are greatly amplified due to moving the cells' internal state across the dynamical phase transition lines. Temporal noise is shown to drive random changes in the cellular polarization direction, which is enhanced during the stick-slip migration mode. These results offer a new framework to explain experimental observations of migrating cells, resulting from noisy switching between underlying deterministic migration modes.

physics.bio-ph↗

arXiv2020

Inverse square Lévy walks are not optimal search strategies for $d\ge 2$

The Lévy hypothesis states that inverse square Lévy walks are optimal search strategies because they maximise the encounter rate with sparse, randomly distributed, replenishable targets. It has served as a theoretical basis to interpret a wealth of experimental data at various scales, from molecular motors to animals looking for resources, putting forward the conclusion that many living organisms perform Lévy walks to explore space because of their optimal efficiency. Here we provide analytically the dependence on target density of the encounter rate of Lévy walks for any space dimension $d$ ; in particular, this scaling is shown to be {\it independent} of the Lévy exponent $α$ for the biologically relevant case $d\ge 2$, which proves that the founding result of the Lévy hypothesis is incorrect. As a consequence, we show that optimizing the encounter rate with respect to $α$ is {\it irrelevant} : it does not change the scaling with density and can lead virtually to {\it any} optimal value of $α$ depending on system dependent modeling choices. The conclusion that observed inverse square Lévy patterns are the result of a common selection process based purely on the kinetics of the search behaviour is therefore unfounded.

cond-mat.stat-mech↗

arXiv2019

Analysis of a non-local and non-linear Fokker-Planck model for cell crawling migration

Cell movement has essential functions in development, immunity and cancer. Various cell migration patterns have been reported and a general rule has recently emerged, the so-called UCSP (Universal Coupling between cell Speed and cell Persistence), [30]. This rule says that cell persistence, which quantifies the straightness of trajectories, is robustly coupled to migration speed. In [30], the advection of polarity cues by a dynamic actin cytoskeleton undergoing flows at the cellular scale was proposed as a first explanation of this universal coupling. Here, following ideas proposed in [30], we present and study a simple model to describe motility initiation in crawling cells. It consists of a non-linear and non-local Fokker-Planck equation, with a coupling involving the trace value on the boundary. In the one-dimensional case we characterize the following behaviours: solutions are global if the mass is below the critical mass, and they can blow-up in finite time above the critical mass. In addition, we prove a quantitative convergence result using relative entropy techniques.

math.AP↗

arXiv2019

Enhanced orientational ordering induced by active yet isotropic bath

Can a bath of isotropic but active particles promote ordering of anisotropic but passive particles? In this paper, we uncover a fluctuation-driven mechanism by which this is possible. Somewhat counter-intuitively, we show that the passive particles tend to be more ordered upon increasing the noise-strength of the active isotropic bath. We first demonstrate this in a general dynamical model for a non-conserved order parameter (model A) coupled to an active isotropic field and then concentrate on two examples, i. a collection of polar rods on a substrate in an active isotropic bath and ii. a passive apolar suspension in a momentum conserved, actively forced but isotropic fluid which is relevant for current research in active systems. Our theory, which is relevant for understanding ordering transitions in out-of-equilibrium systems can be tested in experiments, for instance, by introducing a low concentration of passive rod-like objects in active isotropic fluids and, since it is applicable to any non-conserved dynamical field, may have applications far beyond active matter.

cond-mat.stat-mech↗

arXiv2019

Tunable corrugated patterns in an active gel sheet

Active matter locally converts chemical energy into mechanical work and, for this reason, it provides new mechanisms of pattern formation. In particular, active gels made of protein motors and filaments are far-from-equilibrium systems that exhibit spontaneous flow,[Kruse2004, Voituriez2005] leading to active turbulence in two and three dimensions[Sanchez2012, Kumar2018] and coherent flow in three dimensions[Wu2017] (3D). Although these dynamic flows reveal a characteristic length scale resulting from the interplay between active forcing and passive restoring forces, the observation of static and long-range spatial patterns in active gels has remained elusive. In this work, we demonstrate that a 2D free-standing nematic active gel, formed spontaneously by depletion forces from a 3D solution of kinesin motors and microtubule filaments, actively buckles out-of-plane into a centimeter-sized periodic corrugated sheet that is stable for several days at low activity. Importantly, the corrugations are formed in the absence of flow and their wavelength and stability are controlled by the motor concentration, in agreement with a hydrodynamic theory. At higher activities these patterns are transient with the gel becoming turbulent at longer times. Our results underline the importance of both passive and active forces in shaping active gels and indicate that a static material can be sculpted through an active mechanism.

cond-mat.soft↗

arXiv2018

Signatures of motor susceptibility in the dynamics of a tracer particle in an active gel

We study a model for the motion of a tracer particle inside an active gel, exposing the properties of the van Hove distribution of the particle displacements. Active events of a typical force magnitude give rise to non-Gaussian distributions, having exponential tails or side-peaks. The side-peaks appear when the local bulk elasticity of the gel is large enough and few active sources are dominant. We explain the regimes of the different distributions, and study the structure of the peaks for active sources that are susceptible to the elastic stress that they cause inside the gel. We show how the van Hove distribution is altered by both the duty cycle of the active sources and their susceptibility, and suggest it as a sensitive probe to analyze microrheology data in active systems with restoring elastic forces.

cond-mat.soft↗

arXiv2018

Forces in inhomogeneous open active-particle systems

We study the force that non-interacting point-like active particles apply to a symmetric inert object in the presence of a gradient of activity and particle sources and sinks. We consider two simple patterns of sources and sinks that are common in biological systems. We analytically solve a one dimensional model designed to emulate higher dimensional systems, and study a two dimensional model by numerical simulations. We specify when the particle flux due to the creation and annihilation of particles can act to smooth the density profile that is induced by a gradient in the velocity of the active particles, and find the net resultant force due to both the gradient in activity and the particle flux. These results are compared qualitatively to observations of nuclear motion inside the oocyte, that is driven by a gradient in activity of actin-coated vesicles.

cond-mat.stat-mech↗

arXiv2017

Generalized Archimedes' principle in active fluids

We show how a gradient in the motility properties of non-interacting point-like active particles can cause a pressure gradient that pushes a large inert object. We calculate the force on an object inside a system of active particles with position dependent motion parameters, in one and two dimensions, and show that a modified Archimedes' principle is satisfied. We characterize the system, both in terms of the model parameters and in terms of experimentally measurable quantities: the spatial profiles of the density, velocity and pressure. This theoretical analysis is motivated by recent experiments, which showed that the nucleus of a mouse oocyte (immature egg cell) moves from the cortex to the center due to a gradient of activity of vesicles propelled by molecular motors; it more generally applies to artificial systems of controlled localized activity.

cond-mat.stat-mech↗

arXiv2016

Optimal search strategies of run-and-tumble walks

The run-and-tumble walk, consisting in randomly reoriented ballistic excursions, models phenomena ranging from gas kinetics to bacteria motility. We evaluate the mean time required for this walk to find a fixed target within a 2D or 3D spherical confinement. We find that the mean search time admits a minimum as a function of the mean run duration for various types of boundary conditions and run duration distributions (exponential, power-law, deterministic). Our result stands in sharp contrast to the pure ballistic motion, which is predicted to be the optimal search strategy in the case of Poisson distributed targets.

cond-mat.stat-mech↗