Equilibrium self-assembly and conventional materials processing techniques fall far short of mimicking dynamic self-actuating processes that are commonplace throughout biology. Here, to bridge the gap between living and synthetic matter, we study passive adhesive non-thermal actin fibres immersed in an active microtubule-based fluid. We show that autonomous chaotic flows power non-equilibrium fibre dynamics, thus inducing collisions, generating connections and weaving a membrane-like elastic network. The ensuing active assembly generates a hierarchy of shapes, structures and dynamical processes spanning nanometres to centimetres. Ultimately, it generates an active membrane that exhibits global limit cycles induced by a non-reciprocal coupling between deformations of the elastic membrane and the alignment axis of the nematic active fluid. Our work merges self-assembly with active matter to demonstrate self-processing materials wherein hierarchical life-like structures and dynamics emerge from an initially structureless suspension.

The self-propulsion of +1/2 topological defects is a hallmark of active nematic fluids, where the defects are advected by the flow field that they themselves generate. In this Letter, we propose a minimal model for defect self-propulsion in a nematic active solid: a linear elastic medium with an embedded nematic texture that generates active stress and associated elastic strains. We show that such coupling gives rise to self-propelled +1/2 defects that move relative to the elastic medium by local remodeling of the nematic texture without advection. This mechanism is fundamentally different from the fluid case and can lead to unbinding of defect pairs and stabilization of +1 defects. Our findings might help explain how orientational order, of, for example, muscle fibers, is reconfigured during morphogenesis in solidlike tissues. The proposed mechanism may, for instance, control motility and merging of +1/2 defects, which play a crucial role in setting up the body axis during Hydra regeneration.

After rainfall, pine needles often float on the surface of small puddles. As the water evaporates, they self-assemble into distinct clusters. Motivated by this natural phenomenon, we experimentally investigate the dynamic evolution of synthetic particles as the liquid surface shrinks in area. Our experiments demonstrate the tendency of particles to aggregate, forming distinct clusters as the liquid boundary shrinks. We systematically examine the emergence of these clusters and explore how their sizes and numbers evolve with changes in the packing fraction. We also analyze particle rotation during the process and discuss the formation of the final configuration comprising clusters of various orientations. Complementary numerical simulations demonstrate qualitative agreement with our experimental findings. This study sheds additional light on the self-assembly of macroscopic particles in a dynamically evolving medium.

We study topologically stable defect structures in systems where the defect line classification in three dimensions and associated algebra of interactions (the fundamental group) are governed by the non-Abelian eight-element group, the quaternions Q_{8}. The non-Abelian character of the defect algebra leads to a topological rigidity of bound defect pairs, and trivalent junctions which are the building blocks of multijunction trivalent networks. We realize such structures in laboratory chiral nematics and analyze their behavior analytically, along with numerical modeling.

Biological tissues transform between solid- and liquidlike states in many fundamental physiological events. Recent experimental observations further suggest that in two-dimensional epithelial tissues these solid-liquid transformations can happen via intermediate states akin to the intermediate hexatic phases observed in equilibrium two-dimensional melting. The hexatic phase is characterized by quasi-long-range (power-law) orientational order but no translational order, thus endowing some structure to an otherwise structureless fluid. While it has been shown that hexatic order in tissue models can be induced by motility and thermal fluctuations, the role of cell division and apoptosis (birth and death) has remained poorly understood, despite its fundamental biological role. Here we study the effect of cell division and apoptosis on global hexatic order within the framework of the self-propelled Voronoi model of tissue. Although cell division naively destroys order and active motility facilitates deformations, we show that their combined action drives a liquid-hexatic-liquid transformation as the motility increases. The hexatic phase is accessed by the delicate balance of dislocation defect generation from cell division and the active binding of disclination-antidisclination pairs from motility. We formulate a mean-field model to elucidate this competition between cell division and motility and the consequent development of hexatic order.

Crystalline sheets (e.g., graphene and transition metal dichalcogenides) liberated from a substrate are a paradigm for materials at criticality, because flexural phonons can fluctuate into the third dimension. Although studies of static critical behaviors (e.g., the scale-dependent elastic constants) are plentiful, investigations of dynamics remain limited. Here, we use molecular dynamics to study the time dependence of the midpoint (the height center of mass) of doubly clamped nanoribbons, as prototypical graphene resonators, under a wide range of temperature and strain conditions. By treating the ribbon midpoint as a Brownian particle confined to a nonlinear potential (which assumes a double-well shape beyond the buckling transition), we formulate an effective theory describing the ribbon's transition rate across the two wells and its oscillations inside a given well. We find that, for nanoribbbons compressed above the Euler buckling point and thermalized above a temperature at which the nonlinear effects due to thermal fluctuations become significant, the exponential term (the ratio between energy barrier and temperature) depends only on the geometry but not the temperature, unlike the usual Arrhenius behavior. Moreover, we find that the natural oscillation time for small strain shows a nontrivial scaling tau(o) similar to L-0(z) T (-eta/4), with L-0 being the ribbon length, z = 2 - eta/2 being the dynamic critical exponent, eta = 0.8 being the scaling exponent describing scale-dependent elastic constants, and T being the temperature. These unusual scale- and temperature-dependent dynamics thus exhibit dynamic criticality and could be exploited in the development of graphene-based nanoactuators.

Topological defects play a central role in the physics of many materials, including magnets, superconductors, and liquid crystals. In active fluids, defects become autonomous particles that spontaneously propel from internal active stresses and drive chaotic flows stirring the fluid. The intimate connection between defect textures and active flow suggests that properties of active materials can be engineered by controlling defects, but design principles for their spatiotemporal control remain elusive. Here, we propose a symmetry-based additive strategy for using elementary activity patterns, as active topological tweezers, to create, move, and braid such defects. By combining theory and simulations, we demonstrate how, at the collective level, spatial activity gradients act like electric fields which, when strong enough, induce an inverted topological polarization of defects, akin to a negative susceptibility dielectric. We harness this feature in a dynamic setting to collectively pattern and transport interacting active defects. Our work establishes an additive framework to sculpt flows and manipulate active defects in both space and time, paving the way to design programmable active and living materials for transport, memory, and logic.

Crystalline sheets (e.g., graphene and transition metal dichalcogenides) liberated from a substrate are a paradigm for materials at criticality because flexural phonons can fluctuate into the third dimension. Although studies of static critical behaviors (e.g., the scale-dependent elastic constants) are plentiful, investigations of dynamics remain limited. Here, we use molecular dynamics to study the time dependence of the midpoint (the height center-of-mass) of doubly clamped nanoribbons, as prototypical graphene resonators, under a wide range of temperature and strain conditions. By treating the ribbon midpoint as a Brownian particle confined to a nonlinear potential (which assumes a double-well shape beyond the buckling transition), we formulate an effective theory describing the ribbon's tunneling rate across the two wells and its oscillations inside a given well. We find that, for nanoribbbons compressed above the Euler buckling point and thermalized above a temperature at which the non-linear effects due to thermal fluctuations become significant, the exponential term (the ratio between energy barrier and temperature) depends only on the geometry, but not the temperature, unlike the usual Arrhenius behavior. Moreover, we find that the natural oscillation time for small strain shows a non-trivial scaling $\tau_{\rm o}\sim L_0^{\,z}T^{-\eta/4}$, with $L_0$ being the ribbon length, $z=2-\eta/2$ being the dynamic critical exponent, $\eta=0.8$ being the scaling exponent describing scale-dependent elastic constants, and $T$ being the temperature. These unusual scale- and temperature-dependent dynamics thus exhibit dynamic criticality and could be exploited in the development of graphene-based nanoactuators.

Morphogenesis, the process through which genes generate form, establishes tissue-scale order as a template for constructing the complex shapes of the body plan. The extensive growth required to build these ordered substrates is fuelled by cell proliferation, which, naively, should destroy order. Understanding how active morphogenetic mechanisms couple cellular and mechanical processes to generate order—rather than annihilate it—remains an outstanding question in animal development. We show that cell divisions are the primary drivers of tissue flow, leading to a fourfold orientationally ordered phase. Waves of anisotropic cell proliferation propagate across the embryo with precise patterning. Defects introduced into the nascent lattice by cell divisions are moved out of the tissue bulk towards the boundary by subsequent divisions. Specific cell proliferation rates and orientations enable cell divisions to organize rather than fluidize the tissue. We observe this using live imaging and tissue cartography to analyse the dynamics of fourfold tissue ordering in the trunk segmental ectoderm of the crustacean Parhyale hawaiensis beginning 72 h after egg lay. The result is a robust, active mechanism for generating global orientational order in a non-equilibrium system that sets the stage for the subsequent development of shape and form.

Using the proof of Willmore's conjecture by Marques and Neves, we conjecture that the free energy of tensionless fluid membranes of arbitrary genus has an upper bound. This implies that the average genus of such a membrane, in equilibrium, is finite, regardless of external constraints. We propose that the Gaussian rigidity may be determined by measuring the relative frequencies of large-genus configurations at low temperature.

The vertex model of epithelia describes the apical surface of a tissue as a tiling of polygonal cells, with a mechanical energy governed by deviations in cell shape from preferred, or target, area, A0, and perimeter, P0. The model exhibits a rigidity transition driven by geometric incompatibility as tuned by the target shape index, . For with p*(6) the perimeter of a regular hexagon of unit area, a cell can simultaneously attain both the preferred area and preferred perimeter. As a result, the tissue is in a mechanically soft compatible state, with zero shear and Young's moduli. For p0 < p*(6), it is geometrically impossible for any cell to realize the preferred area and perimeter simultaneously, and the tissue is in an incompatible rigid solid state. Using a mean-field approach, we present a complete analytical calculation of the linear elastic moduli of an ordered vertex model. We analyze a relaxation step that includes non-affine deformations, leading to a softer response than previously reported. The origin of the vanishing shear and Young's moduli in the compatible state is the presence of zero-energy deformations of cell shape. The bulk modulus exhibits a jump discontinuity at the transition and can be lower in the rigid state than in the fluid-like state. The Poisson's ratio can become negative which lowers the bulk and Young's moduli. Our work provides a unified treatment of linear elasticity for the vertex model and demonstrates that this linear response is protocol-dependent.

We study the finite-temperature dynamics of thin elastic sheets in a single-clamped cantilever configuration. This system is known to exhibit a tilt transition at which the preferred mean plane of the sheet shifts from horizontal to a plane above or below the horizontal. The resultant thermally roughened two-state (up/down) system possesses rich dynamics on multiple timescales. In the tilted regime a finite-energy barrier separates the spontaneously chosen up state from the inversion-symmetric down state. Molecular dynamics simulations confirm that, over sufficiently long time, such thermalized elastic sheets transition between the two states, residing in each for a finite dwell time. One might expect that temperature is the primary driver for tilt inversion. We find, instead, that the primary control parameter, at fixed tilt order parameter, is the dimensionless and purely geometrical aspect ratio of the clamped width to the total length of the otherwise-free sheet. Using a combination of an effective mean-field theory and Kramers' theory, we derive the transition rate and examine its asymptotic behavior. At length scales beyond a material-dependent thermal length scale, renormalization of the elastic constants qualitatively modifies the temperature response. In particular, the transition is suppressed by thermal fluctuations, enhancing the robustness of the tilted state. We check and supplement these findings with further molecular dynamics simulations for a range of aspect ratios and temperatures.

Active matter encompasses different nonequilibrium systems in which individual constituents convert energy into non-conservative forces or motion at the microscale. This review provides an elementary introduction to the role of topology in active matter through experimentally relevant examples. Here, the focus lies on topological defects and topologically protected edge modes with an emphasis on the distinctive properties they acquire in active media. These paradigmatic examples represent two physically distinct classes of phenomena whose robustness can be traced to a common mathematical origin: the presence of topological invariants. These invariants are typically integer numbers that cannot be changed by continuous deformations of the relevant order parameters or physical parameters of the underlying medium. We first explain the mechanisms whereby topological defects self propel and proliferate in active nematics, leading to collective states which can be manipulated by geometry and patterning. Possible implications for active microfluidics and biological tissues are presented. We then illustrate how the propagation of waves in active fluids and solids is affected by the presence of topological invariants characterizing their dispersion relations. We discuss the relevance of these ideas for the design of robotic metamaterials and the properties of active granular and colloidal systems. Open theoretical and experimental challenges are presented as future research prospects.

Very thin elastic sheets, even at zero temperature, exhibit nonlinear elastic response by virtue of their dominant bending modes. Their behavior is even richer at finite temperature. Here, we use molecular dynamics to study the vibrations of a thermally fluctuating two-dimensional elastic sheet with one end clamped at its zero-temperature length. We uncover a tilted phase in which the sheet fluctuates about a mean configuration inclined with respect to the horizontal, thus breaking reflection symmetry. We determine the phase behavior as a function of the aspect ratio of the sheet and the temperature. We show that tilt may be viewed as a type of transverse buckling instability induced by clamping coupled to thermal fluctuations and develop an analytic model that predicts the tilted and untilted regions of the phase diagram. Qualitative agreement is found with the molecular dynamics simulations. Unusual response driven by control of purely geometric quantities like the aspect ratio, as opposed to external fields, offers a very rich playground for two-dimensional mechanical metamaterials.

This article summarizes some of the open questions in the field of active matter that have emerged during Active20, a nine-week program held at the Kavli Institute for Theoretical Physics (KITP) in Spring 2020. The article does not provide a review of the field, but rather a personal view of the authors, informed by contributions of all participants, on new directions in active matter research. The topics highlighted include: the ubiquitous occurrence of spontaneous flows and active turbulence and the theoretical and experimental challenges associated with controlling and harnessing such flows; the role of motile topological defects in ordered states of active matter and their possible biological relevance; the emergence of non-reciprocal effective interactions and the role of chirality in active systems and their intriguing connections to non-Hermitian quantum mechanics; the progress towards a formulation of the thermodynamics of active systems thanks to the feedback between theory and experiments; the impact of the active matter framework on our understanding of the emergent mechanics of biological tissue. These seemingly diverse phenomena all stem from the defining property of active matter - assemblies of self-driven entities that individually break time-reversal symmetry and collectively organize in a rich variety of nonequilibrium states.

We study the dynamics of active nematic films on a substrate driven by active flows with or without the incompressible constraint. Through simulations and theoretical analysis, we show that arch patterns are stable in the compressible case, while they become unstable under the incompressibility constraint. For compressible flows at high enough activity, stable arches organize themselves into a smecticlike pattern, which induce an associated global polar ordering of +1/2 nematic defects. By contrast, divergence-free flows give rise to a local nematic order of the +1/2 defects, consisting of antialigned pairs of neighboring defects, as established in previous studies.

We study the active flow around isolated defects and the self-propulsion velocity of + 1 / 2 defects in an active nematic film with both viscous dissipation (with viscosity η ) and frictional damping Γ with a substrate. The interplay between these two dissipation mechanisms is controlled by the hydrodynamic dissipation length ℓ d = η / Γ that screens the flows. For an isolated defect, in the absence of screening from other defects, the size of the shear vorticity around the defect is controlled by the system size R . In the presence of friction that leads to a finite value of ℓ d , the vorticity field decays to zero on the lengthscales larger than ℓ d . We show that the self-propulsion velocity of + 1 / 2 defects grows with R in small systems where R < ℓ d , while in the infinite system limit or when R ≫ ℓ d , it approaches a constant value determined by ℓ d .

Vertex models, such as those used to describe cellular tissue, have an energy controlled by deviations of each cell area and perimeter from target values. The constrained nonlinear relation between area and perimeter leads to new mechanical response. Here we provide a mean-field treatment of a highly simplified model: a uniform network of regular polygons with no topological rearrangements. Since all polygons deform in the same way, we only need to analyze the ground states and the response to deformations of a single polygon (cell). The model exhibits the known transition between a fluid/compatible state, where the cell can accommodate both target area and perimeter, and a rigid/incompatible state. We calculate and measure the mechanical resistance to various deformation protocols and discover that at the onset of rigidity, where a single zero-energy ground state exists, linear elasticity fails to describe the mechanical response to even infinitesimal deformations. In particular, we identify a breakdown of reciprocity expressed via different moduli for compressive and tensile loads, implying nonanalyticity of the energy functional. We give a pictorial representation in configuration space that reveals that the complex elastic response of the vertex model arises from the presence of two distinct sets of reference states (associated with target area and target perimeter). Our results on the critically compatible tissue provide a new route for the design of mechanical metamaterials that violate or extend classical elasticity.

We adapt the Halperin–Mazenko formalism to analyze two-dimensional active nematics coupled to a generic fluid flow. The governing hydrodynamic equations lead to evolution laws for nematic topological defects and their corresponding density fields. We find that ±1/2 defects are propelled by the local fluid flow and by the nematic orientation coupled with the flow shear rate. In the overdamped and compressible limit, we recover the previously obtained active self-propulsion of the +1/2 defects. Non-local hydrodynamic effects are primarily significant for incompressible flows, for which it is not possible to eliminate the fluid velocity in favor of the local defect polarization alone. For the case of two defects with opposite charge, the non-local hydrodynamic interaction is mediated by non-reciprocal pressure-gradient forces. Finally, we derive continuum equations for a defect gas coupled to an arbitrary (compressible or incompressible) fluid flow.