We explore the connection between evolution and large-deviation theory. To do so, we study evolutionary dynamics in which individuals experience mutations, reproduction, and selection using variants of the Moran model. We show that, in the large population size limit, the impact of reproduction and selection amounts to realizing a large-deviation dynamics for the non-interacting random walk in which individuals simply explore the genome landscape due to mutations. This mapping, which holds at all times, allows us to recast transitions in the population genome distribution as dynamical phase transitions, which can then be studied using the toolbox of large-deviation theory. Finally, we show that the mapping extends beyond the class of Moran models.

We study the dynamics of a zero-temperature overdamped tracer in a bath of Brownian particles. As the bath density is increased, numerical simulations show the tracer to transition from an active dynamics, characterized by boundary accumulation and ratchet currents, to an effective equilibrium regime. To account for this analytically, we eliminate the bath degrees of freedom under the assumption of linear coupling to the tracer and show convergence, in the large density limit, to an equilibrium dynamics at the bath temperature. We then develop a perturbation theory to characterize the tracer's departure from equilibrium at large but finite bath densities, revealing an intermediate time-reversible yet non-Boltzmann regime, followed by a fully irreversible one. Finally, we show that when the bath particles are connected as a lattice, mimicking a gel or a soft active solid, the cold tracer drives the entire bath out of equilibrium, leading to a long-ranged suppression of bath fluctuations.

We study the simplest terms that need to be included in active field theories to couple them to external potentials. To do so, we consider active Brownian particles and implement a systematic perturbative expansion in the particle persistence time. The result is a non-trivial coupling between density and potential gradients, which accounts for the nonequilibrium features of active particles in the presence of an external potential, from boundary accumulation to far-field density modulation. We show how the method can be applied to particles interacting via pairwise forces and to spatial modulations of the propulsion speed.

The self-organization of microbial ecosystems involves a large variety of mechanisms, ranging from biochemical signaling to population dynamics. Among these, the role of motility regulation has been little studied, despite the importance of active migration processes. Here we show how weak, random motility regulation suffices to induce complex forms of organization in bacterial mixtures comprising a large number of coexisting strains. First, we simulate microscopic models of run-and-tumble bacteria whose self-propulsion speeds are weakly regulated by the local density of each strain, mimicking the impact of weak, random metabolic interactions. Our simulations reveal that, as the heterogeneity of the interaction network increases, the system undergoes a phase transition leading to the emergence of distinct, spatially segregated communities. To account for these results and assess their robustness, we use random-matrix theory to analyze the hydrodynamic description of the bacterial mixture, obtaining a quantitative agreement with our microscopic simulations. Our results hold for a variety of motility-regulation mechanisms and highlight the need to characterize the role of motility regulation in experimentally relevant situations.

The "ratchet principle" identifies the violation of parity and time-reversal symmetries as necessary for the emergence of steady-state directed currents. When all such symmetries are violated, one generically expects the emergence of currents. We study stochastic systems in the presence of asymmetric fluctuation sources, which violate these symmetries and yet fail to display steady currents. We show that this stems from a hidden conservation law for momentum. For underdamped and overdamped Brownian dynamics, we show that thermal fluctuations cannot power the momentum sources required to sustain directed currents, even when time-reversal symmetry is broken due to an inhomogeneous temperature field. While active Brownian and run-and-tumble particles display interaction-induced directed currents in asymmetric activity landscapes, we show that effective momentum conservation prevents this in Active Ornstein-Uhlenbeck particles: not all inhomogeneous active fluctuations can power transport. For each of the systems considered in this article, we numerically test for the emergence of interaction-induced directed currents. We then characterize time-reversal (a)symmetry in position space using a combination of path-integral and operator methods. When the existence of effective momentum conservation is ruled out, we develop perturbation theories to characterize the onset of interaction-induced directed currents.

We explore the edge flows that emerge at boundaries in nonequilibrium passive and active chiral colloidal fluids. We show that these complex interface currents obey an equation of state that relates their fluxes to bulk observables. For confined fluids, the edge flux is given by the average odd stress in the fluid. In phase-separated systems, the flux along the interface is given by the jump of the odd stress across the interface. We then use the equation of state to reveal, and contrast, the microscopic origins of the edge currents in passive and active systems.

We study correlation functions in generic non-equilibrium mixtures, including multi-temperature systems and non-reciprocal field theories. The corresponding linear theory is short-ranged, and nonlinearities are irrelevant in the renormalization-group sense. Nonetheless, we find that these nonlinearities generate long-ranged three-point correlations in the isotropic disordered phase. Our analytical predictions, which are based on a phenomenological theory, are confirmed by numerical simulations of Brownian colloids in contact with thermal baths at different temperatures. Dangerously irrelevant nonlinearities in non-equilibrium mixtures thus offer a new route to long-range correlations, supporting the hypothesis that such correlations are not the exception but the rule out of equilibrium.

We study the stability of the ordered phase of compressible polar flocks against the nucleation of counter-propagating droplets, using a combination of analytical theory, microscopic and hydrodynamic simulations. For discrete-symmetry flocks, such droplets are known to always grow and propagate, making the ordered phase metastable. We explain how, on the contrary, continuous symmetry can stabilize the ordered phase at small enough noise by destabilizing the leading edge of growing droplets. Flocking models with continuous symmetries thus have a lower critical dimension than their discrete-symmetry counterparts, in contrast to equilibrium physics.

We study the dynamics of weakly deformed interfaces separating two stable phases, starting from the fluctuating hydrodynamics of the phase-separating fields. Using a well-chosen definition for the interface and the dynamical-action formalism to represent path probabilities, we derive the linear relaxation of the interface and the fluctuations around it for a large class of models. Our method applies to equilibrium dynamics, where it recovers and complements existing results, but also extends to their non-equilibrium counterparts. We explain how non-linear terms can be systematically computed and illustrate their derivations in the case of (active) model A. We highlight the danger of a popular ansatz used to derive interface dynamics, which was rigorously established in equilibrium but is uncontrolled for active field theories.

The Young-Dupr & eacute; equation is a cornerstone of the theory of capillary and wetting phenomena, which governs the shape of droplets adsorbed on surfaces in equilibrium. However, many living and synthetic materials, such as swarming bacteria and active colloids, are composed of self-propelled particles that are inherently out of thermal equilibrium. The description of the wetting of surfaces by such active fluids thus requires a new framework. Here we develop an analogue to the Young-Dupr & eacute; equation for systems made of self-propelled particles. A key step is to define the liquid-gas surface tension of active fluids as the force exerted along the interface, which we show from first principles to be negative, even when active materials separate into stable liquid and gas phases. Our active Young-Dupr & eacute; equation explains why partial wetting appears in simulations where the surface tensions do not balance and reveals the underlying feedback mechanism: the interface is stable only because of steady flows, which are themselves generated by the parity symmetry-breaking interface. Unlike in passive fluids, where the droplets are scale-free, this feedback loop selects the sizes and shapes of adsorbed droplets in active materials. Our results outline a framework for understanding how active matter wets surfaces.

We study the dynamics of a zero-temperature particle interacting linearly with a bath of hot Brownian particles. Starting with the most general model of a linearly-coupled bath, we eliminate the bath degrees of freedom exactly to map the tracer dynamics onto a generalized Langevin equation, allowing for an arbitrary external potential on the tracer. We apply this result to determine the fate of a tracer connected by springs to N identical bath particles or inserted within a harmonic chain of hot particles. In the former "fully-connected" case, we find the tracer to transition between an effective equilibrium regime at large N and an FDT-violating regime at finite N, while in the latter "loop" model the tracer never satisfies an FDT. We then study the fully-connected model perturbatively for large but finite N, demonstrating signatures of irreversibility such as ratchet currents, non-Boltzmann statistics, and positive entropy production. Finally, we specialize to harmonic external potentials on the tracer, allowing us to exactly solve the dynamics of both the tracer and the bath for an arbitrary linear model. We apply our findings to show that a cold tracer in a hot lattice suppresses the fluctuations of the lattice in a long-ranged manner, and we generalize this result to linear elastic field theories.

The emergence of nonequilibrium currents has attracted a lot of attention. The "ratchet principle" identifies the violation of any form of parity and time-reversal symmetries as the necessary conditions for the emergence of such currents. Here we study active and passive systems with asymmetric spatially varying fluctuation sources that nevertheless remain flux free. For non-interacting systems, we show that a hidden time-reversal symmetry forbids the emergence of steady currents. At higher densities, pairwise forces break this symmetry but an emergent conservation law for the momentum field may nevertheless prevent steady currents. We show how the presence of this conservation law can be tested analytically and characterize the onset of ratchet currents in its absence. Our results show that the ratchet principle should be refined to also preclude bulk momentum conservation.

Unlike biological active matter that constantly adapt to their environment, the motors of synthetic active particles are typically agnostic to their surroundings and merely operate at constant force. Here, we design colloidal active rods capable of modulating their inner activity in response to crowding, thereby enforcing a primitive form of quorum sensing interactions. Through experiments, simulations, and theory we elucidate the impact of these interactions on the phase behavior of isotropic active matter. We demonstrate that, when conditioned to density, motility regulation can either lead to an absorbing phase transition, where all particles freeze their dynamics, or to atypical phase separation, where flat interfaces supporting a net pressure drop are in mechanical equilibrium. Fully active and fully arrested particles can then form heterogeneous patterns ruled by the competition between quorum sensing and mechanical interactions. Beyond the specifics of motile colloids, we expect our findings to apply broadly to adaptive active matter assembled from living or synthetic units.

Long-range interactions are ubiquitous in nature, where they are mediated by diffusive fields at the cellular scale or by visual cues for groups of animals. Short-range forces, which are paradigmatic in physics, can thus often be neglected when modeling the collective behaviors of biological systems induced by mediated interactions. However, when self-organization leads to the emergence of dense phases, we show that excluded-volume interactions play an important and versatile role. We consider assemblies of active particles that undergo either condensation or phase-separation due to motility regulation and show that short-range repulsive forces can induce opposite effects. When motility regulation triggers an absorbing phase transition, such as a chemotactic collapse, repulsive forces opposes the formation of condensates and stabilize the coexistence between finite-density phases. In contrast, when motility regulation induces liquid-gas coexistence, repulsive forces can, counterintuitively, lead to a significant increase in the liquid density.

We present a detailed derivation of the Langevin dynamics obeyed by a massive rigid body immersed in a chiral active bath. We show how the antisymmetric nature of the noise leads to an unusual relationship between the Langevin equation describing stochastic trajectories and the Fokker-Planck equation governing the time-evolution of the probability density. The chirality of the bath endows the object dynamics with odd diffusivity, odd mobility, and rotational ratchet effects that depend on the object symmetries. For rotationally-symmetric objects, we show that a hidden time-reversal symmetry leads to separate effective equilibrium descriptions for the translational and rotational degrees of freedom. Finally, starting from the bath dynamics, we construct a multipole expansion to quadrupolar order that allows predicting the far-field current and density modulation induced by the object on the bath.

The "ratchet principle" asserts that non-equilibrium systems which violate parity symmetry generically exhibit steady-state currents. As recently shown, there are exceptions to this principle, due to the existence of hidden time-reversal symmetry or bulk momentum conservation. For underdamped and overdamped Brownian dynamics, we show how thermal fluctuations cannot power the momentum sources required to sustain steady ratchet currents, even when time-reversal symmetry is broken due to an inhomogeneous temperature field. While Active Brownian and Run-and-Tumble particles display interaction-induced ratchet currents in asymmetric activity landscapes, we show that this is not the case for Active Ornstein-Uhlenbeck particles: not all inhomogeneous active fluctuations lead to net momentum sources. For each of the systems considered in this article, we numerically test for the emergence of interaction-induced ratchet currents. We then characterize time-reversal (a)symmetry in position space using a combination of path-integral and operator methods. When the existence of effective momentum conservation is ruled out, we develop perturbation theories to characterize the onset of interaction-induced currents.

When submerged in a chiral active bath, a passive object becomes a spinning ratchet imbued with odd transport properties. We present the most general Langevin dynamics for a rigid body in a chiral active bath, in the adiabatic limit of large object mass. For rotationally symmetric objects, odd diffusion and odd mobility are connected by an Einstein relation, that we show numerically to break down outside the adiabatic limit. As the object symmetry decreases, its dynamics becomes increasingly irreversible: a massive disk exhibits an effective equilibrium dynamics, while a rod admits distinct translational and rotational temperatures, and a wedge is fully irreversible.Conversely, this departure from equilibrium can be read in universal far-field currents and density modulations of the bath, which we measure numerically and derive analytically.

We present a detailed derivation of the Langevin dynamics obeyed by a massive rigid body immersed in a chiral active bath. We show how the antisymmetric nature of the noise leads to an unusual relationship between the Langevin equation describing stochastic trajectories and the Fokker-Planck equation governing the time evolution of the probability density. The chirality of the bath endows the object dynamics with odd diffusivity, odd mobility, and rotational ratchet effects that depend on the object symmetries. For rotationally symmetric objects, we show that a hidden time-reversal symmetry leads to separate effective equilibrium descriptions for the translational and rotational degrees of freedom. Finally, starting from the bath dynamics, we construct a multipole expansion to quadrupolar order that allows predicting the far-field current and density modulation induced by the object on the bath.

In this article we derive and test the fluctuating hydrodynamic description of active particles interacting via taxis and quorum sensing, both for mono-disperse systems and for mixtures of co-existing species of active particles. We compute the average steady-state density profile in the presence of spatial motility regulation, as well as the structure factor and intermediate scattering function for interacting systems. By comparing our predictions to microscopic numerical simulations, we show that our fluctuating hydrodynamics correctly predicts the large-scale static and dynamical properties of the system. We also discuss how the theory breaks down when structures emerge at scales smaller or comparable to the persistence length of the particles. When the density field is the unique hydrodynamic mode of the system, we show that active Brownian particles, run-and-tumble particles and active Ornstein-Uhlenbeck particles, interacting via quorum-sensing or chemotactic interactions, display undistinguishable large-scale properties. This form of universality implies an interesting robustness of the predicted physics but also that large-scale observations of patterns are insufficient to assess their microscopic origins. In particular, our results predict that chemotaxis-induced and motility-induced phase separation should share strong qualitative similarities at the macroscopic scale.

The nature of the transition to collective motion in assemblies of aligning self-propelled particles remains a long-standing matter of debate. In this article, we focus on dry active matter and show that weak fluctuations suffice to generically turn second-order mean-field transitions into a 'discontinuous' coexistence scenario. Our theory shows how fluctuations induce a density-dependence of the polar-field mass, even when this effect is absent at mean-field level. In turn, this dependency on density triggers a feedback loop between ordering and advection that ultimately leads to an inhomogeneous transition to collective motion and the emergence of inhomogeneous travelling bands. Importantly, we show that such a fluctuation-induced first order transition is present in both metric models, in which particles align with neighbors within a finite distance, and in 'topological' ones, in which alignment is based on more complex constructions of neighbor sets. We compute analytically the noise-induced renormalization of the polar-field mass using stochastic calculus, which we further back up by a one-loop field-theoretical analysis. Finally, we confirm our analytical predictions by numerical simulations of fluctuating hydrodynamics as well as of topological particle models with either k-nearest neighbors or Voronoi alignment.