Raphaël Voituriez’s research while affiliated with Theoretical physics of condensed matter and other places

In this chapter, we consider the problem of a non-Markovian random walker (displaying memory effects) searching for a target. We review an approach that links the first passage statistics to the properties of trajectories followed by the random walker in the future of the first passage time. This approach holds in one and higher spatial dimensions, when the dynamics in the vicinity of the target is Gaussian, and it is applied to three paradigmatic target search problems: the search for a target in confinement, the search for a rarely reached configuration (rare event kinetics), or the search for a target in infinite space, for processes featuring stationary increments or transient aging. The theory gives access to the mean first passage time (when it exists) or to the behavior of the survival probability at long times, and agrees with the available exact results obtained perturbatively for examples of weakly non-Markovian processes. This general approach reveals that the characterization of the non-equilibrium state of the system at the instant of first passage is key to derive first-passage kinetics, and provides a new methodology, via the analysis of trajectories after the first-passage, to make it quantitative.

The persistence exponent, which characterises the long-time decay of the survival probability of stochastic processes in the presence of an absorbing target, plays a key role in quantifying the dynamics of fluctuating systems. Determining this exponent for non-Markovian processes is known to be a difficult task, and exact results remain scarce despite sustained efforts. In this Letter, we consider the fundamental class of self-interacting random walks (SIRWs), which display long-range memory effects that result from the interaction of the random walker at time t with the territory already visited at earlier times $t'<t$. We compute exactly the persistence exponent for all physically relevant SIRWs. As a byproduct, we also determine the splitting probability of these processes. Besides their intrinsic theoretical interest, these results provide a quantitative characterization of the exploration process of SIRWs, which are involved in fields as diverse as foraging theory, cell biology, and machine learning.

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, with 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. Significance statement During embryonic development and wound healing, epithelial cells usually display in-plane polarity over large spatial scales and move coherently. However, over years, most in vitro studies have examined the fluid-like chaotic dynamics of epithelial cells, in which collective cellular flows self-organize into recurring transient vortices and jets similar to those observed in classical fluid turbulence. Little is known about the large-scale coherent dynamics of epithelial cells. We demonstrate that such coherent motions are not simply turbulent-like flows with larger correlation lengths, but a new mode of collective motion with a solid-like behavior, accompanied by an emergent global order, scale-free correlations, anomalous density fluctuations and propagating Goldstone modes. Our work suggests that such a collective motion of epithelial cells falls outside the scope of traditional active fluids, which may shed new light on the current studies of collective cell migration as well as active matter physics.

Self-interacting random walks (SIRWs) show long-range memory effects that result from the interaction of the random walker at time t with the territory already visited at earlier times t^{'}<t. This class of non-Markovian random walks has applications in contexts as diverse as foraging theory, the behavior of living cells, and even machine learning. Despite this importance and numerous theoretical efforts, the propagator, which is the distribution of the walker's position and arguably the most fundamental quantity to characterize the process, has so far remained out of reach for all but a single class of SIRW. Here we fill this gap and provide an exact and explicit expression for the propagator of two important universality classes of SIRWs, namely, the once-reinforced random walk and the polynomially self-repelling walk. These results give access to key observables, such as the diffusion coefficient, which so far had not been determined. We also uncover an inherently non-Markovian mechanism that tends to drive the walker away from its starting point.

Flocking is a prime example of how robust collective behavior can emerge from simple interaction rules. The flocking transition has been studied extensively since the inception of the original Vicsek model. Here, we introduce a novel self-propelled particle model with quenched disorder in the pairwise alignment interaction couplings akin to a spin glass model. We show that the presence of quenched disorder can promote (rather than destroy) the emergence of global polar order. In particular, we show that our model can display a flocking phase even when the majority of the interaction couplings are anti-aligning. Activity is the key ingredient to reduce frustration in the system as it allows local particle clustering combined with self-organization of the particles to favor neighborhoods with strong cooperative interactions.

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.

The Kramers escape problem is a paradigmatic model for the kinetics of rare events, which are usually characterized by Arrhenius law. So far, analytical approaches have failed to capture the kinetics of rare events in the important case of non-Markovian processes with long-term memory, as occurs in the context of reactions involving proteins, long polymers, or strongly viscoelastic fluids. Here, based on a minimal model of non-Markovian Gaussian process with long-term memory, we determine quantitatively the mean FPT to a rare configuration and provide its asymptotics in the limit of a large energy barrier E. Our analysis unveils a correction to Arrhenius law, induced by long-term memory, which we determine analytically. This correction, which we show can be quantitatively significant, takes the form of a second effective energy barrier E′<E$E^{\prime} \, < \, E$ and captures the dependence of rare event kinetics on initial conditions, which is a hallmark of long-term memory. Altogether, our results quantify the impact of long-term memory on rare event kinetics, beyond Arrhenius law.

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.

Biological surfaces, such as developing epithelial tissues, exhibit in-plane polar or nematic order and can be strongly curved. Recently, integer (+1) topological defects have been identified as morphogenetic hotspots in living systems. Yet, while +1 defects in active matter on flat surfaces are well-understood, the general principles governing curved active defects remain unknown. Here, we study the dynamics of integer defects in an extensile or contractile polar fluid on two types of morphogenetically-relevant substrates : (1) a cylinder terminated by a spherical cap, and (2) a bump on an otherwise flat surface. Because the Frank elastic energy on a curved surface generically induces a coupling to $\textit{deviatoric}$ curvature, $\mathcal{D}$ (difference between squared principal curvatures), a +1 defect is induced on both surface types. We find that $\mathcal{D}$ leads to surprising effects including localization of orientation gradients and active flows, and particularly for contractility, to hysteresis and bistability between quiescent and flowing defect states.