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Active matter

Active matter refers to a class of non-equilibrium physical systems composed of numerous interacting units, such as particles or agents, that are self-propelled and convert free energy from their environment or internal sources into sustained, directed motion and mechanical work.^[1] These systems operate far from thermodynamic equilibrium, consuming energy to drive persistent activity that leads to emergent collective behaviors not observed in passive matter. Unlike equilibrium systems, active matter exhibits broken time-reversal symmetry and can generate long-range correlations and ordered phases through local energy inputs. In biological contexts, active matter manifests in systems like swarming bacteria (e.g., Escherichia coli exhibiting run-and-tumble motility), molecular motors such as kinesin walking along microtubules, and larger-scale phenomena including bird flocks and fish schools, where individual self-propulsion and interactions yield complex patterns like milling or vortex formation.^[1] Synthetic realizations include active colloids, such as Janus particles that propel via asymmetric chemical reactions or phoretic effects, and light- or magnetically driven microswimmers designed in laboratories to mimic biological motility. These examples span scales from nanometers (e.g., cytoskeletal filaments) to meters (e.g., animal groups), highlighting the universality of active principles across biology and engineered materials. The physics of active matter draws from statistical mechanics, hydrodynamics, and non-equilibrium thermodynamics to explain phenomena like phase transitions to ordered states (e.g., flocking in the Vicsek model), giant number fluctuations in dilute suspensions, and instabilities in active gels where contractile forces lead to spontaneous flow. Key challenges include developing a thermodynamic framework for energy dissipation and entropy production in these systems, as well as understanding how activity couples to passive degrees of freedom like orientation or shape deformations. Research has revealed that active matter can violate detailed balance and produce novel states, such as active nematics with topological defects driving spontaneous motion.^[1] Emerging over the past two decades at the intersection of soft matter physics, biophysics, and materials science, active matter has spurred applications in micro-robotics for targeted drug delivery, soft robotics mimicking biological swarms, and understanding cellular processes like tissue morphogenesis. Ongoing studies explore active matter under extreme conditions, such as microgravity, to probe universal principles of self-organization.^[1] This field continues to grow, with theoretical models informing experimental designs and revealing how local activity generates global order in living and artificial systems.

Definition and Fundamentals

Definition

Active matter refers to collections of self-propelled agents that consume energy from their environment to generate systematic motion or mechanical forces, thereby maintaining the system in a state far from thermodynamic equilibrium.^[2] These agents, which can include biological entities like bacteria or synthetic particles, operate through internal mechanisms that drive persistent, directed movement without relying on external concentration gradients.^[2] In contrast to passive matter, which obeys detailed balance and time-reversal symmetry in equilibrium conditions, active matter inherently breaks these symmetries due to continuous local energy dissipation, enabling phenomena such as spontaneous collective organization and nontrivial steady-state currents. This nonequilibrium character arises from the unidirectional conversion of energy into work at the individual agent level, distinguishing active systems from passive ones where motion ceases without external driving.^[2] Active matter manifests across a wide range of scales, from molecular levels—such as motor proteins that propel along cytoskeletal filaments—to macroscopic assemblies like flocks of birds or schools of fish.^[2] At these scales, the agents convert environmental energy sources into mechanical work; in biological contexts, this often involves chemical energy from ATP hydrolysis, while synthetic realizations may harness external inputs like light or electric fields to induce propulsion.^[2]^[3]

Key Characteristics

Active matter systems are distinguished by the persistent, directed motion of their constituent agents, which arises from internal energy dissipation rather than external forces. This persistence is quantified by the persistence length l_p = v_0 \tau, where v_0 is the characteristic self-propulsion speed and \tau is the reorientation time, representing the distance an agent travels before significantly changing direction. In models of active Brownian particles, this length scale emerges from the competition between directed propulsion and rotational diffusion, leading to superdiffusive trajectories over intermediate times. A central metric for activity in these systems is the Péclet number Pe = v_0 L / D, which compares advective transport due to self-propulsion over a system size L to diffusive spreading governed by the diffusion coefficient D. High Pe values indicate regimes where directed motion dominates, enabling collective phenomena absent in passive systems. This parameter is particularly relevant in dilute suspensions, where it controls the onset of instabilities and clustering. Unlike equilibrium fluids, active matter exhibits giant number fluctuations, where the relative density fluctuations scale as \langle (\delta n)^2 \rangle / \langle n \rangle^2 \sim N^{2/d - 1} in d dimensions (e.g., independent of subsystem size N in 2D), contrasting with the equilibrium scaling \sim 1/N.^[2] These anomalous fluctuations, first predicted in models of oriented active suspensions, arise from the coupling between density and orientational order, amplified by activity. Experimental observations in bacterial suspensions and synthetic colloids confirm this enhanced variability, signaling nonequilibrium long-range correlations. Activity also drives the emergence of orientational order and correlations, even in low dimensions where thermal fluctuations would destroy it in equilibrium systems. In two dimensions, polar flocks develop long-range orientational order, as described by the Vicsek model, due to velocity alignment interactions sustained by persistent motion.^[4] Nematic order similarly persists, with correlations extending over macroscopic scales, fostering collective flows and defects.

Historical Development

Early Biological Observations

The earliest scientific observations of active matter phenomena in biological systems date back to the late 19th century, when botanist Wilhelm Pfeffer described directed movements of microorganisms in response to chemical gradients in 1884, coining the term "chemotaxis" based on experiments with bacteria and sperm cells.^[5] Pfeffer's work highlighted how these self-propelled motions allowed organisms to navigate environments without external forces, laying foundational empirical evidence for nonequilibrium dynamics in living systems. In the mid-20th century, these observations were extended to specific bacterial behaviors, with Julius Adler's 1966 studies on Escherichia coli demonstrating chemotactic responses to nutrients like sugars and amino acids, revealing a bias in random motion toward favorable conditions. Further insights into bacterial motility emerged from three-dimensional tracking experiments, which visualized the "run-and-tumble" pattern in E. coli, where cells alternate straight swimming runs with random reorientations via tumbling, enabling efficient exploration and chemotaxis.^[6] Concurrently, observations of intracellular dynamics revealed active contractions driving cell shape changes, as seen in amoeboid movement studied in the early 20th century. Microscopic examinations of protozoa like Amoeba proteus showed rhythmic contractions of cytoplasmic fibrils, suggesting a contractile apparatus akin to muscle, with later biochemical identification of actin-like proteins in non-muscle cells, such as in the slime mold Physarum polycephalum, confirming myosin-actin interactions powered by ATP hydrolysis. These findings underscored self-driven cytoskeletal remodeling as a key mechanism for cellular motility, dissipating metabolic energy to maintain far-from-equilibrium states. Collective behaviors in multicellular organisms provided additional early examples of coordinated active motion without central control. In fish, C.M. Breder's 1976 analysis of schooling patterns described how groups maintain parallel orientation and spacing through local interactions, optimizing hydrodynamic efficiency and predator avoidance in species like herring and sardines.^[7] Similarly, insect swarms, such as those of midges (Chironomidae), were observed in the mid-20th century to exhibit synchronized hovering and directional shifts, driven by visual cues and wind, as documented in field studies of lekking behaviors where males form dynamic aerial displays. These pre-1990s empirical accounts recognized chemotaxis and phototaxis—light-directed motility in algae like Euglena—as inherently active processes fueled by continuous energy input from metabolism, distinguishing them from passive diffusion and foreshadowing the dissipative nature of biological activity.

Emergence of Theoretical Models

The emergence of theoretical models for active matter in the mid-1990s marked a shift from qualitative biological descriptions to quantitative physics, inspired by observations of collective animal behaviors such as bird flocking and fish schooling. A foundational contribution was the Vicsek model, introduced in 1995, which simulates self-propelled particles that align their velocities with neighbors within a fixed radius while subject to random noise, revealing a phase transition to collective ordered motion above a critical noise threshold.^[8] This agent-based approach demonstrated how local alignment rules could lead to global coherence without centralized control, laying the groundwork for studying emergent order in driven systems. Concurrently, John Toner and Yuhai Tu developed a hydrodynamic theory for flocking, starting with a 1995 paper that proposed a dynamical XY model for polar-ordered flocks and extended it through 1998 to derive continuum equations capturing long-range order and giant fluctuations in two dimensions. Their framework includes the momentum equation

\partial_t \mathbf{v} + (\mathbf{v} \cdot \nabla) \mathbf{v} = -\nabla P + \nu \nabla^2 \mathbf{v} + \mathbf{f},

where \mathbf{v} is the velocity field, P is pressure, \nu is viscosity, and \mathbf{f} represents active and noise terms, highlighting anomalous scaling behaviors distinct from equilibrium systems. In the early 2000s, Sriram Ramaswamy and collaborators advanced the field by exploring active nematics, systems of rod-like particles with orientational order but no polar axis, showing in 2003 that such models on substrates exhibit giant density fluctuations and long-time correlations due to activity-induced instabilities.^[9] Ramaswamy's work also introduced the classification of active matter into "dry" systems, where momentum is not conserved due to substrate friction, and "wet" systems, which include hydrodynamic interactions in a fluid medium, providing a conceptual dichotomy for modeling diverse active phenomena.^[10] By 2013, M. C. Marchetti and colleagues synthesized these developments into a unified hydrodynamic framework for soft active matter, integrating microscopic origins with continuum descriptions to predict generic instabilities, phase transitions, and nonequilibrium patterns across biological and synthetic systems.^[10] This review emphasized the role of symmetry and activity in driving collective behaviors, establishing active matter as a distinct branch of nonequilibrium statistical physics.

Theoretical Frameworks

Microscopic Models

Microscopic models in active matter focus on the dynamics of individual self-propelled particles, capturing their stochastic motion and interactions through discrete agent-based rules or differential equations, which enable simulations of emergent phenomena at the particle level. These approaches emphasize bottom-up descriptions, where active forces and orientational persistence drive behaviors without deriving coarse-grained fields. Seminal formulations include run-and-tumble dynamics for abrupt reorientations and persistent random walks for smoother turning, often extended with alignment rules for collective effects.^[11] Run-and-tumble particles (RTPs) represent a key microscopic model for organisms exhibiting ballistic motion interrupted by random reorientations, such as flagellated bacteria. In this framework, each particle i propels at constant speed v_0 along its orientation \hat{\mathbf{u}}_i during a "run" phase, with position updating as \dot{\mathbf{r}}_i = v_0 \hat{\mathbf{u}}_i. Tumbles occur stochastically at rate $1/\tau, instantaneously randomizing the direction \hat{\mathbf{u}}_i uniformly on the unit circle (in 2D) or sphere (in 3D). This model captures persistence over run length v_0 \tau and has been pivotal in studying motility-induced phase separation, where interactions like volume exclusion lead to clustering despite no explicit attraction.^[12]^[13] Active Brownian particles (ABPs) provide another foundational microscopic description, modeling overdamped swimmers with continuous rotational diffusion, suitable for colloidal or eukaryotic systems. The dynamics follow the Langevin equations \dot{\mathbf{r}} = v_0 \hat{\mathbf{u}} + \sqrt{2D} \boldsymbol{\xi} for translational motion, where \boldsymbol{\xi} is Gaussian white noise with \langle \boldsymbol{\xi}(t) \boldsymbol{\xi}(t') \rangle = \mathbf{I} \delta(t-t'), and \dot{\theta} = \sqrt{2 D_r} \eta for orientation, with \eta similarly Gaussian noise and D_r the rotational diffusivity. This yields a persistent random walk with persistence time $1/D_r and effective diffusion D_{\text{eff}} = D + v_0^2 / (2 D_r) at long times. ABPs are widely used to explore density-dependent slowdowns and giant number fluctuations in interacting suspensions.^[14]^[13] Extensions of the Vicsek model introduce alignment interactions to microscopic particle dynamics, promoting flocking from local velocity averaging. In the original metric version, each particle updates its direction toward the average of neighbors within radius R, plus noise: \hat{\mathbf{u}}_i(t + \Delta t) = \text{normalize} \left( \sum_{j \in N_i} \hat{\mathbf{u}}_j(t) \right) + \boldsymbol{\eta}_i, where N_i are particles within Euclidean distance R of i. Topological variants replace metric distance with fixed-neighbor counts, such as the first C nearest neighbors via Voronoi shells, enhancing robustness to density variations as observed in bird flocks. These models often incorporate repulsion or attraction potentials, like Lennard-Jones for exclusion or Morse for binding, to stabilize clusters without hydrodynamic effects.^[8]^[15]^[16] Simulating these microscopic models adapts molecular dynamics techniques to include active propulsion as non-conservative forces, with event-driven or velocity-Verlet integrators handling overdamped limits via Brownian thermostats. Packages like LAMMPS extend standard MD by adding self-propulsion terms and custom noise for orientations, enabling efficient handling of millions of particles. Computational challenges arise from long-time correlations in persistent motion, requiring large system sizes (N > 10^4) and extended runs (t > 10^3 \tau) to resolve slow diffusion and avoid finite-size artifacts in order parameters like polarization P = \left| \frac{1}{N} \sum_i \hat{\mathbf{u}}_i \right|.^[17]^[11]

Hydrodynamic and Continuum Approaches

Hydrodynamic and continuum approaches to active matter provide coarse-grained descriptions that capture collective behaviors emerging from underlying microscopic dynamics, such as self-propulsion and interactions among agents. These field-theoretic models treat the system as continuous fields—density, velocity, or orientation—governed by partial differential equations derived via symmetry considerations or systematic coarse-graining from particle-based theories. Unlike equilibrium hydrodynamics, these nonequilibrium equations incorporate activity-driven terms that break time-reversal symmetry, leading to novel instabilities and long-range correlations. Such approaches are essential for understanding large-scale phenomena in both dry (friction-dominated) and wet (momentum-conserving) active systems.^[18] A cornerstone of these theories for polar active matter, where agents align and move collectively, is the Toner-Tu framework, originally developed for flocking behaviors like bird schools. The model consists of coupled equations for the velocity field \mathbf{v} and density \rho, with the velocity dynamics featuring nonlinear advection and damping terms that stabilize long-range orientational order even in two dimensions, defying the Mermin-Wagner theorem through nonequilibrium effects. The key equation for the velocity is

\partial_t \mathbf{v} + \lambda (\mathbf{v} \cdot \nabla) \mathbf{v} = \Gamma \nabla^2 \mathbf{v} - \alpha \mathbf{v} + \beta |\mathbf{v}|^2 \mathbf{v} + \dots,

where \lambda governs nonlinear advection, \Gamma is a diffusion coefficient, \alpha and \beta control linear instability and saturation, and the dots represent additional nonlinearities, pressure gradients, and noise. This form arises from coarse-graining microscopic alignment rules and self-propulsion, predicting ordered phases with mean velocity \langle \mathbf{v} \rangle \neq 0. Renormalization group analysis of these equations yields anomalous scaling exponents, such as the dynamical exponent z = 6/5, anisotropy exponent ζ = 3/5, and roughness exponent χ = -1/5 in two dimensions, ensuring the stability of long-range order.^[19]^[18] For scalar active matter without intrinsic polarity, such as run-and-tumble bacteria or active colloids, continuum theories extend equilibrium models like Cahn-Hilliard to include activity. The Active Model B+ is a paradigmatic example, describing density \phi dynamics via a conserved current with an active contribution that mimics effective attractions from density-dependent motility slowdowns. The governing equation is

\partial_t \phi = \nabla^2 \left( \frac{\delta F}{\delta \phi} - \zeta \phi \right),

where F[\phi] is the equilibrium free energy (e.g., Ginzburg-Landau form), and \zeta > 0 parameterizes activity, driving phase separation beyond equilibrium spinodals. This model, derived from microscopic theories of self-propelled particles with density-dependent speeds, predicts motility-induced phase separation (MIPS), where dense clusters form without explicit attractions, stabilized by activity. Microscopic origins lie in agent-based simulations of persistent random walks, but the continuum limit reveals universal coarsening dynamics.^[20] In apolar active matter, such as suspensions of rod-like swimmers, active nematics emerge with orientational order described by the nematic tensor Q_{ij}. The hydrodynamic equations couple Q to flow via an active stress contribution \sigma^a_{ij} = \zeta Q_{ij}, where \zeta sets the activity strength (negative for extensile systems like microtubules, positive for contractile). This term enters the force balance, generating spontaneous flows that destabilize uniform states: extensile nematics favor splay instabilities (director divergence), while contractile ones promote bend deformations (director curl), leading to chaotic flows and defect proliferation at long wavelengths. These instabilities, analyzed linearly, grow as \sigma \sim q^2 for small wavevector q, contrasting passive liquid crystals.^[21] A hallmark of these continuum theories is the presence of giant fluctuations, far exceeding equilibrium Poisson statistics. In polar flocks and active nematics, density fluctuations scale as \delta N \sim N^{1/2 + \alpha}, with \alpha > 0 from renormalization group flows that couple orientation to density. For two-dimensional active nematics on substrates, \alpha = 1/4, yielding \delta N \sim N^{3/4}, while polar flocks exhibit \alpha = 3/10. These exponents, computed perturbatively to one loop, highlight how activity amplifies correlations, observable in experiments on bacterial vortices and synthetic rollers.^[18]^[22]

Experimental Realizations

Biological Systems

Biological systems exemplify active matter through self-sustaining, energy-dissipating processes that drive non-equilibrium dynamics at multiple scales, from intracellular structures to large animal groups. At the cellular level, the cytoskeleton functions as an active gel, where actin filaments and microtubules interact with molecular motors to generate persistent stresses and flows. Myosin motors, in particular, bind to actin filaments and slide them relative to one another, producing contractile forces that reorganize the network into coherent patterns such as asters—star-like radial assemblies—and vortices, which emerge from the interplay of motor-induced stresses and filament polarity.^[23]^[24]^[25] In microbial communities, bacterial swarms demonstrate active matter behaviors through density-dependent motility that leads to clustering analogous to motility-induced phase separation (MIPS). In experiments with self-propelled bacterial-like rods in the 2000s, such as those modeling Escherichia coli and other species, collisions and volume exclusion cause particles to slow in dense regions, fostering aggregation into stable clusters without attractive forces. These MIPS-like phenomena have been observed in confined E. coli suspensions, where collective motion results in phase-separated domains, highlighting how bacterial self-propulsion drives emergent spatial organization.^[26]^[27] Tissue dynamics further illustrate active fluid behavior in multicellular assemblies, particularly in epithelial sheets where cells maintain cohesion while undergoing flows driven by proliferation and death. Cell division introduces local expansions that fluidize the tissue, while apoptosis creates voids that contribute to directed flows, effectively rendering the sheet a viscoelastic active fluid with a relaxation timescale governed by these rates. Seminal models show that this activity enables tissue spreading and reshaping, as seen in embryonic development and wound healing, where division and apoptosis balance to sustain coherent monolayer flows without external forcing.^[28] At the macroscopic scale, collectives of higher organisms exhibit flocking and herding as quintessential active matter phenomena, characterized by long-range order and velocity alignment. In bird flocks, such as those of European starlings (Sturnus vulgaris), real-time stereo-tracking of thousands of individuals reveals scale-free correlations in positions and velocities, with birds maintaining topological interactions with about seven neighbors regardless of density, enabling rapid, coherent maneuvers. Similarly, herding in mammals like goats demonstrates analogous collective motion, where individuals align velocities through local interactions, as quantified by high-resolution tracking in natural groups, underscoring shared principles of self-organization across scales.^[29]

Synthetic Systems

Synthetic active matter encompasses engineered systems composed of non-living components that mimic the self-propulsion and collective dynamics observed in biological active matter, enabling precise control over activity levels, interactions, and emergent behaviors through external fields or chemical triggers. These systems are typically fabricated at micro- or nanoscales using colloidal particles, emulsions, or macroscopic robots, allowing researchers to tune parameters such as speed, directionality, and density to study fundamental principles of nonequilibrium physics. Unlike biological systems, synthetic realizations emphasize scalability and programmability, facilitating experiments that probe phase transitions and pattern formation under controlled conditions. A foundational example of synthetic active matter is colloidal microswimmers, particularly Janus particles, which are bifacial colloids with one side coated in a catalytic material to generate autonomous propulsion. In a seminal demonstration, striped nanorods consisting of platinum and gold segments, approximately 370 nm in diameter and 1 μm long, were shown to move at speeds up to 10 body lengths per second in hydrogen peroxide solutions through catalytic decomposition that produces oxygen bubbles or chemical gradients driving phoretic motion. This self-propulsion arises from asymmetric surface reactions, where the platinum catalyzes H₂O₂ decomposition into O₂ and H₂O, creating a local concentration gradient that propels the particle away from the reactive side. Subsequent refinements have explored shape effects on propulsion direction, confirming that geometric asymmetry enhances directional control in these catalytic swimmers. Quincke rotors represent another class of synthetic active particles, where dielectric colloids in a uniform DC electric field undergo spontaneous rotation due to electrohydrodynamic instabilities, converting rotational motion into translation near surfaces. These rollers, typically micrometer-sized particles, exhibit tunable speeds up to several body lengths per second and form active nematic phases when densely packed, displaying defects and spontaneous flows characteristic of nonequilibrium liquid crystals. In bulk fluids, Quincke rotation powers active particles without contact, enabling studies of chaotic dynamics governed by Lorenz-like equations. Complementing these, active emulsions involve self-propelled oil-in-water droplets driven by interfacial tensions or chemical reactions, such as Belousov-Zhabotinsky oscillations, which induce Marangoni flows for propulsion. These droplets, often stabilized by surfactants, exhibit chemotactic behaviors and phase separation in dense suspensions, with collective dynamics tunable via topology and surfactant concentration, as seen in systems where double emulsions form 3D active structures with programmable flow patterns. Macroscopic synthetic active matter is exemplified by robotic swarms like the Kilobots, low-cost (about $20 each) vibration-driven robots that demonstrate programmable collective behaviors at scales of thousands of units. Developed in the 2010s, these 3.3 cm disc-shaped robots use infrared communication to follow simple rules for self-assembly into shapes or flocking patterns, achieving synchronization and decision-making akin to natural swarms without central control. In one large-scale experiment, 1,024 Kilobots autonomously formed complex two-dimensional structures, such as stars or letters, by local gradient-based algorithms, highlighting scalability for testing emergent phenomena like consensus formation. Recent advances up to 2025 have pushed synthetic active matter toward hybrid nanoscale systems with enhanced responsiveness. Light-activated DNA origami swimmers integrate folded DNA nanostructures with photoresponsive elements, such as azobenzene-modified strands, to enable on-demand propulsion via UV-induced conformational changes that drive cyclic motion along tracks. These systems, often combined with catalytic or magnetic components, achieve directed swimming at speeds of micrometers per second under visible light, offering reconfigurability for studying active transport. Similarly, 3D-printed active metamaterials incorporate stimuli-responsive polymers or ferrofluids into lattice structures, allowing tunable activity through magnetic or thermal fields; for instance, microlattice shells filled with magnetorheological fluid exhibit stiffness modulation over orders of magnitude, enabling programmable mechanical responses in active assemblies.

Emergent Behaviors

Collective Motion and Flocking

Collective motion in active matter refers to the emergence of coherent, ordered group dynamics where individual agents align their orientations and velocities, leading to large-scale patterns such as flocks or schools. This phenomenon is driven by local alignment rules in the presence of noise and interactions, resulting in a nonequilibrium phase transition from disordered to ordered states. Unlike passive systems, active matter exhibits spontaneous symmetry breaking and long-range order even in the absence of external fields, as self-propulsion breaks time-reversal symmetry.^[8] A paradigmatic model for flocking transitions is the Vicsek model, where self-propelled particles move at constant speed and periodically align their directions with neighbors within a fixed interaction radius, perturbed by random noise. The order parameter quantifying the degree of alignment is defined as \psi = \left| \sum_{i=1}^N \hat{\mathbf{u}}_i / N \right|, where \hat{\mathbf{u}}_i is the unit velocity vector of particle i and N is the total number of particles; \psi \approx 0 in the disordered phase and \psi \approx 1 in the ordered flocking phase. The model undergoes a continuous phase transition at a critical noise threshold, with the infinite-system noise strength \eta_c(\infty) \approx 2.9 (for uniform noise in [-\eta/2, \eta/2] and density \rho = 0.4), below which ordered motion emerges.^[8]^[30] In biological flocks, such as starling murmurations, interactions are often topological rather than metric, meaning agents respond to a fixed number of nearest neighbors (typically around 7) irrespective of distance, rather than all within a fixed radius. This topological coupling enhances flock robustness against perturbations, maintaining cohesion and order more effectively than metric rules, as evidenced by field studies tracking over 3000 birds.^[15] Milling and vortex states represent confined variants of collective motion, where agents form rotating patterns due to boundary effects and alignment. In circular confinements, bacterial suspensions can stabilize into coherent spiral vortices, with cells circulating en masse around the perimeter, as observed in experiments with Bacillus subtilis. These states arise from hydrodynamic interactions and chiral biases, contrasting with unconfined linear flocks.^[31] Anomalous correlations in flocking manifest as power-law decay in velocity fields, \langle \mathbf{v}(\mathbf{r}) \cdot \mathbf{v}(0) \rangle \sim r^{-\alpha} with \alpha < d (where d is dimension), leading to giant fluctuations far exceeding equilibrium diffusion predictions. This nonequilibrium feature, differing from exponential decay in passive fluids, arises from the coupling of orientation and density in hydrodynamic descriptions of flocks.

Phase Separation and Transitions

One prominent example of activity-driven structural changes in active matter is motility-induced phase separation (MIPS), where self-propelled particles with purely repulsive interactions spontaneously demix into dense and dilute phases despite lacking any attractive forces. This phenomenon arises from a feedback mechanism: in dense regions, particle collisions reduce motility, leading to an effective attraction that accumulates particles and further slows their motion. In models of active Brownian particles (ABPs), the phase diagram features a binodal line separating coexisting dense and dilute phases, as well as a spinodal line delineating unstable regions where fluctuations amplify rapidly, analogous to equilibrium liquid-gas transitions but driven out-of-equilibrium. At sufficiently high densities, active matter can exhibit active glass transitions characterized by arrested dynamics, where particles become trapped in disordered configurations with dramatically slowed diffusion and relaxation times. Unlike passive glasses, these transitions occur without thermal quenching and can persist even at low effective temperatures due to persistent self-propulsion.^[32] In active gels—cross-linked networks of motile elements such as microtubules and motors—these arrested states develop a yield stress, below which the material behaves as a solid, while above it flows viscously, enabling tunable rheological properties. For elongated or rod-like active particles, such as bacterial flagella or synthetic swimmers, activity promotes orientational ordering into nematic and smectic phases, where particles spontaneously align parallel or form layered structures. These phases emerge from hydrodynamic interactions and self-propulsion that favor alignment, but they are prone to instabilities, including spontaneous tumbling or bending modes that disrupt order and generate chaotic flows. Experimental observations have confirmed these transitions in diverse systems. In synthetic setups, MIPS has been realized with colloidal rollers propelled by optical traps or magnetic fields, where dense clusters form and coarsen over time in two dimensions during the 2020s. Similarly, bacterial suspensions, such as Vibrio cholerae or Escherichia coli, exhibit MIPS-like clustering in confined geometries, driven by steric hindrance to motility in crowded regions.

Applications and Implications

Biological and Biomedical Uses

Active matter principles have been instrumental in elucidating the dynamics of wound healing, where collective cell migration drives tissue repair through active stresses that generate flows in epithelial sheets. In this process, epithelial cells exhibit polar ordering, modeled as active polar fluids where contractile stresses and substrate friction propel monolayer expansion to close wounds. For instance, analytical models treating the epithelial layer as a viscous polar fluid with active contractility predict spreading velocities dependent on gap size and purse-string mechanisms at the wound edge, aligning with experimental observations of re-epithelialization kinetics. Similarly, in morphogenesis, active nematic phases emerge in developing tissues, with topological defects facilitating cell rearrangements and apoptosis to shape tissue structures. These models highlight how active stresses, akin to those in polar active gels, coordinate multicellular flows during embryonic development.^[33]^[34]^[35] In cancer metastasis, active matter concepts frame collective invasion as an unjamming transition, where tumor cells shift from a solid-like jammed state to a fluid-like motile phase, enabling dissemination from the primary site. This transition, driven by changes in cell shape, motility, and adhesion, occurs at non-zero densities and is modulated by epithelial-mesenchymal transition (EMT), allowing clusters of cancer cells to invade the extracellular matrix collectively. Experimental studies on breast cancer spheroids reveal phase diagrams mapping jammed cores to unjammed peripheries, with ECM stiffness dictating invasion modes—collective fluid-like in low-density matrices and single-cell gas-like in others. Such insights suggest therapeutic strategies targeting jamming regulators, like adhesion molecules or motility pathways, to prevent unjamming and metastasis, potentially broadening drug interventions beyond traditional biochemical targets.^[36]^[37]^[38] Self-propelled microrobots, embodying active matter propulsion, offer promising avenues for targeted drug delivery in biomedical applications, harnessing biological motors for precise navigation in vivo. Biohybrid systems, such as sperm-hybrid micromotors, integrate motile sperm cells with magnetic microstructures to load and release drugs like doxorubicin at tumor sites, achieving high encapsulation efficiency (up to 98%) and localized cytotoxicity against cancer spheroids with minimal leakage. These devices exploit sperm's natural chemotaxis and biocompatibility for gynecological applications, with magnetic guidance enabling controlled release upon target contact. In the 2020s, advances have progressed in preclinical models of biohybrid microrobots, demonstrating enhanced tumor penetration and reduced systemic toxicity.^[39]^[40]^[41] Recent applications of active matter in organoids leverage these principles to model diseases through emergent multicellular behaviors in self-assembled tissues. Brain organoids, modeled as active gels, exhibit activity-induced instabilities like folding, driven by contractile stresses that mimic cortical gyrification and reveal pathological dynamics in neurodevelopmental disorders. These models capture non-equilibrium phase separations and polar flows, enabling simulation of disease states. By integrating patient-derived stem cells, active matter frameworks in organoids facilitate high-throughput screening of emergent behaviors, advancing personalized disease modeling and therapeutic discovery. As of 2025, light-controlled programming of biological active matter has enabled micrometer-scale fluid flow fields for transport and separation in organoid-like systems.^[42]^[43]

Engineering and Materials Science

In engineering and materials science, active matter principles have been harnessed to develop soft robotics systems capable of collective behaviors without centralized control. Swarming microbots, inspired by biological collectives, enable tasks such as environmental sensing through distributed exploration and adaptation. For instance, extensions of Harvard's Kilobot platforms in the 2020s have led to link-bots, which are 3D-printed, centimeter-scale particles connected in V-shaped chains that exhibit emergent locomotion via steric interactions and vibrations. These systems demonstrate adaptive morphologies, allowing chains to reconfigure for navigation through obstacles or object transport, with applications in real-time environmental monitoring where swarms self-organize to map chemical or physical gradients.^[44] Active metamaterials represent another key advancement, integrating active elements to achieve dynamic, programmable mechanical properties. These structures incorporate embedded oscillators, such as non-reciprocally coupled bistable units, to drive shape-shifting behaviors and enable tunable responses to external stimuli. For example, mechanical metamaterials with oscillator arrays can propagate unidirectional solitons, facilitating reconfigurable stiffness and deformation patterns that mimic living tissues. Such designs allow for shape morphing through reversible polymorphic transformations, where the material shifts between states like expansion or contraction in response to thermal or mechanical cues, offering programmable mechanics for applications in adaptive devices.^[45]^[46] Energy harvesting from active matter has focused on active colloids for microfluidic applications, where self-generated temperature gradients are converted into directed fluid flow. Anisotropic thermophoretic active colloids serve as autonomous pumps in microchannels, eliminating the need for external power sources. These systems achieve efficient small-scale fluid transport by leveraging thermophoresis, with demonstrated pumping rates suitable for lab-on-a-chip devices. By 2025, optimizations in colloid design have improved conversion efficiencies of thermal energy to mechanical work, enabling sustained flows in confined environments for precise delivery in sensing or actuation tasks.^[47] Scalability remains a core challenge in deploying active matter systems, addressed through 3D printing of active composites that embed responsive elements like shape-memory polymers. Direct 4D printing techniques allow fabrication of high-resolution structures (down to 50 μm) with built-in strain, enabling rapid transformation and reprogrammability without post-processing. This approach overcomes limitations in traditional manufacturing by producing scalable, multi-shape composites that maintain active functionality at larger scales, such as in deployable robotics or metamaterial arrays. Recent reviews highlight how such methods mitigate issues like material inhomogeneity, facilitating industrial translation of active matter into robust engineering solutions.^[48]^[49]

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Jul 19, 2013 · This review summarizes theoretical progress in the field of active matter, placing it in the context of recent experiments.
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[1910.02528] Computational models for active matter - arXiv
Oct 6, 2019 · This paper reviews computational models for active matter, from microscopic to macroscopic scales, addressing challenges like out-of- ...Missing: simulation | Show results with:simulation
[12]
Motility-induced phase separation and coarsening in active matter
Most of this work will focus on the case of run-and-tumble and active Brownian particles in the absence of solvent-mediated hydrodynamic interactions – we will ...Missing: seminal | Show results with:seminal<|separator|>
[13]
Active Brownian particles and run-and-tumble particles separate ...
Nov 23, 2016 · In this article we have explored the behavior of two important and widespread types of active particles inside different mazes. Namely, non- ...
[14]
Active Brownian Particles. From Individual to Collective Stochastic ...
Feb 11, 2012 · We review theoretical models of individual motility as well as collective dynamics and pattern formation of active particles.
[15]
Interaction ruling animal collective behavior depends on topological ...
Interaction ruling animal collective behavior depends on topological rather than metric distance: Evidence from a field study. M. Ballerini, N. Cabibbo, R.
[16]
Relevance of Metric-Free Interactions in Flocking Phenomena
We introduce a “topological Vicsek model” in which constant-speed particles align their velocity with that of others forming their first Voronoi shell. We show ...
[17]
Molecular Dynamics Simulations of Active Matter using LAMMPS
Feb 20, 2021 · LAMMPS is a widely popular classical Molecular Dynamics package. It was designed for materials modeling but it is well prepared for simulations in Soft Matter.
[18]
Active gels as a description of the actin-myosin cytoskeleton - PMC
This short review presents a qualitative introduction to the hydrodynamic theory of active polar gels and its applications to the mechanics of the cytoskeleton.Missing: seminal | Show results with:seminal
[19]
Active contractility in actomyosin networks - PNAS
Actomyosin networks generate contractile forces through the activity of myosin motors, which themselves assemble into bipolar minifilaments that generate ...Active Contractility In... · Sign Up For Pnas Alerts · Main Findings<|control11|><|separator|>
[20]
Asters, Vortices, and Rotating Spirals in Active Gels of Polar Filaments
Feb 20, 2004 · We develop a general theory for active viscoelastic materials made of polar filaments. This theory is motivated by the dynamics of the cytoskeleton.Missing: seminal | Show results with:seminal
[21]
Nonequilibrium clustering of self-propelled rods | Phys. Rev. E
We study collective motion in a two-dimensional model of active, self-propelled rods interacting through volume exclusion.
[22]
Collective motion in an active suspension of Escherichia coli bacteria
Feb 13, 2014 · We investigate experimentally the emergence of collective motion in the bulk of an active suspension of Escherichia coli bacteria.
[23]
Fluidization of tissues by cell division and apoptosis - PMC
We show that the tissue effectively behaves as a viscoelastic fluid with a relaxation time set by the rates of division and apoptosis.Missing: sheets seminal
[24]
Dynamics of collective motion across time and species - Journals
Feb 20, 2023 · We study the collective motion of four species: shoals of stickleback fish (Gasterosteus aculeatus), flocks of homing pigeons (Columba livia), a herd of goats.
[25]
https://link.aps.org/doi/10.1103/PhysRevLett.92.078101
[26]
Confinement Stabilizes a Bacterial Suspension into a Spiral Vortex
Jun 24, 2013 · We investigate here the influence of global confinement and surface curvature on collective motion by studying the flow and orientational order ...Missing: milling seminal paper
[27]
Nonequilibrium glass transitions in driven and active matter - arXiv
Feb 20, 2013 · This paper studies nonequilibrium glass transitions in driven and active matter, showing dynamic arrest can occur, and that effective thermal ...
[28]
Dense active matter model of motion patterns in confluent cell ...
Mar 16, 2020 · Wound healing revised: a novel reepithelialization mechanism ... Generic theory of active polar gels: a paradigm for cytoskeletal dynamics.
[29]
The analytical solution to the migration of an epithelial monolayer ...
Aug 29, 2022 · Such tissue is usually used to mimic the wound healing process in Virto. We model the epithelial sheet as a layer of active viscous polar fluid.
[30]
Topology-guided polar ordering of collective cell migration - Science
Apr 17, 2024 · A prominent example of active matter in biology comes from studies demonstrating the presence of active nematic order, a class of active matter ...
[31]
Jamming transitions in cancer - PMC - PubMed Central - NIH
Cellular jamming differs from jamming in inert matter in two aspects: first, cells are active particles, and second, the jamming transition occurs at a packing ...Missing: drug | Show results with:drug
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https://arxiv.org/abs/1302.4868
[33]
EMT-related cell-matrix interactions are linked to states of cell ...
Dec 20, 2024 · Epithelial-to-mesenchymal transitions (EMT) and unjamming transitions provide two distinct pathways for cancer cells to become invasive, ...
[34]
Sperm-Hybrid Micromotor for Targeted Drug Delivery | ACS Nano
Dec 4, 2017 · A sperm-driven micromotor is presented as a targeted drug delivery system, which is appealing to potentially treat diseases in the female reproductive tract.
[35]
IRONSperm: Sperm-templated soft magnetic microrobots - Science
Jul 8, 2020 · Sperm micromotors have also been shown to deliver anticancer drugs locally to cancer spheroids (29). On the other hand, another proposed method ...
[36]
Biohybrid microrobots locally and actively deliver drug-loaded ... - NIH
Jun 12, 2024 · Here, we show a microrobot approach using motile algae for localized delivery of drug-loaded nanoparticles to address lung metastasis challenges ...Missing: matter 2020s<|separator|>
[37]
Activity-induced instabilities of brain organoids - PMC - NIH
We present an active matter model system for describing the experimentally observed folds on brain organoids. ... Modeling development and disease with organoids.
[38]
Emergent functional dynamics of link-bots | Science Advances
May 9, 2025 · We investigate how simple steric interaction constraints between active individuals produce a versatile and functional system using the link-bot.
[39]
Non-reciprocal topological solitons in active metamaterials - arXiv
Dec 6, 2023 · Here we introduce a local driving mechanism for solitons that accelerates both solitons and antisolitons in the same direction instead: non-reciprocal driving.
[40]
Shape morphing mechanical metamaterials through reversible ...
Feb 9, 2022 · We introduce a multifunctional shape-morphing material with reversible and rapid polymorphic reconfigurability.
[41]
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[42]
Direct 4D printing via active composite materials | Science Advances
Apr 12, 2017 · We describe an approach to print composite polymers in high-resolution three-dimensional (3D) architectures that can be rapidly transformed ...
[43]
3D printing of active mechanical metamaterials: A critical review
This study offered a promising approach for designing programmable load plateaus and developing novel metamaterials with tunable force–displacement responses ...