Machine learning interpretable models of cell mechanics from protein images.

Cellular form and function emerge from complex mechanochemical systems within the cytoplasm. Currently, no systematic strategy exists to infer large-scale physical properties of a cell from its molecular components. This is an obstacle to understanding processes such as cell adhesion and migration. Here, we develop a data-driven modeling pipeline to learn the mechanical behavior of adherent cells. We first train neural networks to predict cellular forces from images of cytoskeletal proteins. Strikingly, experimental images of a single focal adhesion (FA) protein, such as zyxin, are sufficient to predict forces and can generalize to unseen biological regimes. Using this observation, we develop two approaches-one constrained by physics and the other agnostic-to construct data-driven continuum models of cellular forces. Both reveal how cellular forces are encoded by two distinct length scales. Beyond adherent cell mechanics, our work serves as a case study for integrating neural networks into predictive models for cell biology.

Pattern formation by turbulent cascades.

Fully developed turbulence is a universal and scale-invariant chaotic state characterized by an energy cascade from large to small scales at which the cascade is eventually arrested by dissipation1-6. Here we show how to harness these seemingly structureless turbulent cascades to generate patterns. Pattern formation entails a process of wavelength selection, which can usually be traced to the linear instability of a homogeneous state7. By contrast, the mechanism we propose here is fully nonlinear. It is triggered by the non-dissipative arrest of turbulent cascades: energy piles up at an intermediate scale, which is neither the system size nor the smallest scales at which energy is usually dissipated. Using a combination of theory and large-scale simulations, we show that the tunable wavelength of these cascade-induced patterns can be set by a non-dissipative transport coefficient called odd viscosity, ubiquitous in chiral fluids ranging from bioactive to quantum systems8-12. Odd viscosity, which acts as a scale-dependent Coriolis-like force, leads to a two-dimensionalization of the flow at small scales, in contrast with rotating fluids in which a two-dimensionalization occurs at large scales4. Apart from odd viscosity fluids, we discuss how cascade-induced patterns can arise in natural systems, including atmospheric flows13-19, stellar plasma such as the solar wind20-22, or the pulverization and coagulation of objects or droplets in which mass rather than energy cascades23-25.

Non-reciprocal phase transitions.

Out of equilibrium, a lack of reciprocity is the rule rather than the exception. Non-reciprocity occurs, for instance, in active matter1-6, non-equilibrium systems7-9, networks of neurons10,11, social groups with conformist and contrarian members12, directional interface growth phenomena13-15 and metamaterials16-20. Although wave propagation in non-reciprocal media has recently been closely studied1,16-20, less is known about the consequences of non-reciprocity on the collective behaviour of many-body systems. Here we show that non-reciprocity leads to time-dependent phases in which spontaneously broken continuous symmetries are dynamically restored. We illustrate this mechanism with simple robotic demonstrations. The resulting phase transitions are controlled by spectral singularities called exceptional points21. We describe the emergence of these phases using insights from bifurcation theory22,23 and non-Hermitian quantum mechanics24,25. Our approach captures non-reciprocal generalizations of three archetypal classes of self-organization out of equilibrium: synchronization, flocking and pattern formation. Collective phenomena in these systems range from active time-(quasi)crystals to exceptional-point-enforced pattern formation and hysteresis. Our work lays the foundation for a general theory of critical phenomena in systems whose dynamics is not governed by an optimization principle.

Spiking at the edge: Excitability at interfaces in reaction-diffusion systems.

Excitable media, ranging from bioelectric tissues and chemical oscillators to forest fires and competing populations, are nonlinear, spatially extended systems capable of spiking. Most investigations of excitable media consider situations where the amplifying and suppressing forces necessary for spiking coexist at every point in space. In this case, spikes arise due to local bistabilities, which require a fine-tuned ratio between local amplification and suppression strengths. But, in nature and engineered systems, these forces can be segregated in space, forming structures like interfaces and boundaries. Here, we show how boundaries can generate and protect spiking when the reacting components can spread out: Even arbitrarily weak diffusion can cause spiking at the edge between two non-excitable media. This edge spiking arises due to a global bistability, which can occur even if amplification and suppression strengths do not allow spiking when mixed. We analytically derive a spiking phase diagram that depends on two parameters: i) the ratio between the system size and the characteristic diffusive length-scale and ii) the ratio between the amplification and suppression strengths. Our analysis explains recent experimental observations of action potentials at the interface between two non-excitable bioelectric tissues. Beyond electrophysiology, we highlight how edge spiking emerges in predator-prey dynamics and in oscillating chemical reactions. Our findings provide a theoretical blueprint for a class of interfacial excitations in reaction-diffusion systems, with potential implications for spatially controlled chemical reactions, nonlinear waveguides and neuromorphic computation, as well as spiking instabilities, such as cardiac arrhythmias, that naturally occur in heterogeneous biological media.

Dualities and non-Abelian mechanics.

Dualities are mathematical mappings that reveal links between apparently unrelated systems in virtually every branch of physics1-8. Systems mapped onto themselves by a duality transformation are called self-dual and exhibit remarkable properties, as exemplified by the scale invariance of an Ising magnet at the critical point. Here we show how dualities can enhance the symmetries of a dynamical matrix (or Hamiltonian), enabling the design of metamaterials with emergent properties that escape a standard group theory analysis. As an illustration, we consider twisted kagome lattices9-15, reconfigurable mechanical structures that change shape by means of a collapse mechanism9. We observe that pairs of distinct configurations along the mechanism exhibit the same vibrational spectrum and related elastic moduli. We show that these puzzling properties arise from a duality between pairs of configurations on either side of a mechanical critical point. The critical point corresponds to a self-dual structure with isotropic elasticity even in the absence of spatial symmetries and a twofold-degenerate spectrum over the entire Brillouin zone. The spectral degeneracy originates from a version of Kramers' theorem16,17 in which fermionic time-reversal invariance is replaced by a hidden symmetry emerging at the self-dual point. The normal modes of the self-dual systems exhibit non-Abelian geometric phases18,19 that affect the semiclassical propagation of wavepackets20, leading to non-commuting mechanical responses. Our results hold promise for holonomic computation21 and mechanical spintronics by allowing on-the-fly manipulation of synthetic spins carried by phonons.

Motor crosslinking augments elasticity in active nematics.

In active materials, uncoordinated internal stresses lead to emergent long-range flows. An understanding of how the behavior of active materials depends on mesoscopic (hydrodynamic) parameters is developing, but there remains a gap in knowledge concerning how hydrodynamic parameters depend on the properties of microscopic elements. In this work, we combine experiments and multiscale modeling to relate the structure and dynamics of active nematics composed of biopolymer filaments and molecular motors to their microscopic properties, in particular motor processivity, speed, and valency. We show that crosslinking of filaments by both motors and passive crosslinkers not only augments the contributions to nematic elasticity from excluded volume effects but dominates them. By altering motor kinetics we show that a competition between motor speed and crosslinking results in a nonmonotonic dependence of nematic flow on motor speed. By modulating passive filament crosslinking we show that energy transfer into nematic flow is in large part dictated by crosslinking. Thus motor proteins both generate activity and contribute to nematic elasticity. Our results provide new insights for rationally engineering active materials.

Liquid-crystal-based topological photonics.

Liquid crystals are complex fluids that allow exquisite control of light propagation thanks to their orientational order and optical anisotropy. Inspired by recent advances in liquid-crystal photo-patterning technology, we propose a soft-matter platform for assembling topological photonic materials that holds promise for protected unidirectional waveguides, sensors, and lasers. Crucial to our approach is to use spatial variations in the orientation of the nematic liquid-crystal molecules to emulate the time modulations needed in a so-called Floquet topological insulator. The varying orientation of the nematic director introduces a geometric phase that rotates the local optical axes. In conjunction with suitably designed structural properties, this geometric phase leads to the creation of topologically protected states of light. We propose and analyze in detail soft photonic realizations of two iconic topological systems: a Su-Schrieffer-Heeger chain and a Chern insulator. The use of soft building blocks potentially allows for reconfigurable systems that exploit the interplay between topological states of light and the underlying responsive medium.

Machine learning active-nematic hydrodynamics.

Hydrodynamic theories effectively describe many-body systems out of equilibrium in terms of a few macroscopic parameters. However, such parameters are difficult to determine from microscopic information. Seldom is this challenge more apparent than in active matter, where the hydrodynamic parameters are in fact fields that encode the distribution of energy-injecting microscopic components. Here, we use active nematics to demonstrate that neural networks can map out the spatiotemporal variation of multiple hydrodynamic parameters and forecast the chaotic dynamics of these systems. We analyze biofilament/molecular-motor experiments with microtubule/kinesin and actin/myosin complexes as computer vision problems. Our algorithms can determine how activity and elastic moduli change as a function of space and time, as well as adenosine triphosphate (ATP) or motor concentration. The only input needed is the orientation of the biofilaments and not the coupled velocity field which is harder to access in experiments. We can also forecast the evolution of these chaotic many-body systems solely from image sequences of their past using a combination of autoencoders and recurrent neural networks with residual architecture. In realistic experimental setups for which the initial conditions are not perfectly known, our physics-inspired machine-learning algorithms can surpass deterministic simulations. Our study paves the way for artificial-intelligence characterization and control of coupled chaotic fields in diverse physical and biological systems, even in the absence of knowledge of the underlying dynamics.

Spatiotemporal control of liquid crystal structure and dynamics through activity patterning.

Active materials are capable of converting free energy into mechanical work to produce autonomous motion, and exhibit striking collective dynamics that biology relies on for essential functions. Controlling those dynamics and transport in synthetic systems has been particularly challenging. Here, we introduce the concept of spatially structured activity as a means of controlling and manipulating transport in active nematic liquid crystals consisting of actin filaments and light-sensitive myosin motors. Simulations and experiments are used to demonstrate that topological defects can be generated at will and then constrained to move along specified trajectories by inducing local stresses in an otherwise passive material. These results provide a foundation for the design of autonomous and reconfigurable microfluidic systems where transport is controlled by modulating activity with light.

Active Viscoelasticity of Odd Materials.

The mechanical response of active media ranging from biological gels to living tissues is governed by a subtle interplay between viscosity and elasticity. We generalize the canonical Kelvin-Voigt and Maxwell models to active viscoelastic media that break both parity and time-reversal symmetries. The resulting continuum theories exhibit viscous and elastic tensors that are both antisymmetric, or odd, under exchange of pairs of indices. We analyze how these parity violating viscoelastic coefficients determine the relaxation mechanisms and wave-propagation properties of odd materials.

Topological Defects in Solids with Odd Elasticity.

Crystallography typically studies collections of point particles whose interaction forces are the gradient of a potential. Lifting this assumption generically gives rise in the continuum limit to a form of elasticity with additional moduli known as odd elasticity. We show that such odd elastic moduli modify the strain induced by topological defects and their interactions, even reversing the stability of, otherwise, bound dislocation pairs. Beyond continuum theory, isolated dislocations can self propel via microscopic work cycles active at their cores that compete with conventional Peach-Koehler forces caused, for example, by an ambient torque density. We perform molecular dynamics simulations isolating active plastic processes and discuss their experimental relevance to solids composed of spinning particles, vortexlike objects, and robotic metamaterials.

Erratum: Active Viscoelasticity of Odd Materials [Phys. Rev. Lett. 126, 138001 (2021)].

This corrects the article DOI: 10.1103/PhysRevLett.126.138001.

Localizing softness and stress along loops in 3D topological metamaterials.

Topological states can be used to control the mechanical properties of a material along an edge or around a localized defect. The rigidity of elastic networks is characterized by a topological invariant called the polarization; materials with a well-defined uniform polarization display a dramatic range of edge softness depending on the orientation of the polarization relative to the terminating surface. However, in all 3D mechanical metamaterials proposed to date, the topological modes are mixed with bulk soft modes, which organize themselves in Weyl loops. Here, we report the design of a 3D topological metamaterial without Weyl lines and with a uniform polarization that leads to an asymmetry between the number of soft modes on opposing surfaces. We then use this construction to localize topological soft modes in interior regions of the material by including defect lines-dislocation loops-that are unique to three dimensions. We derive a general formula that relates the difference in the number of soft modes and states of self-stress localized along the dislocation loop to the handedness of the vector triad formed by the lattice polarization, Burgers vector, and dislocation-line direction. Our findings suggest a strategy for preprogramming failure and softness localized along lines in 3D, while avoiding extended soft Weyl modes.

Soft self-assembly of Weyl materials for light and sound.

Soft materials can self-assemble into highly structured phases that replicate at the mesoscopic scale the symmetry of atomic crystals. As such, they offer an unparalleled platform to design mesostructured materials for light and sound. Here, we present a bottom-up approach based on self-assembly to engineer 3D photonic and phononic crystals with topologically protected Weyl points. In addition to angular and frequency selectivity of their bulk optical response, Weyl materials are endowed with topological surface states, which allow for the existence of one-way channels, even in the presence of time-reversal invariance. Using a combination of group-theoretical methods and numerical simulations, we identify the general symmetry constraints that a self-assembled structure has to satisfy to host Weyl points and describe how to achieve such constraints using a symmetry-driven pipeline for self-assembled material design and discovery. We illustrate our general approach using block copolymer self-assembly as a model system.

Symmetries and Dualities in the Theory of Elasticity.

Microscopic symmetries impose strong constraints on the elasticity of a crystalline solid. In addition to the usual spatial symmetries captured by the tensorial character of the elastic tensor, hidden nonspatial symmetries can occur microscopically in special classes of mechanical structures. Examples of such nonspatial symmetries occur in families of mechanical metamaterials where a duality transformation relates pairs of different configurations. We show on general grounds how the existence of nonspatial symmetries further constrains the elastic tensor, reducing the number of independent moduli. In systems exhibiting a duality transformation, the resulting constraints on the number of moduli are particularly stringent at the self-dual point but persist even away from it, in a way reminiscent of critical phenomena.

Non-Hermitian Band Topology and Skin Modes in Active Elastic Media.

Solids built out of active components can exhibit nonreciprocal elastic coefficients that give rise to non-Hermitian wave phenomena. Here, we investigate non-Hermitian effects present at the boundary of two-dimensional active elastic media obeying two general assumptions: their microscopic forces conserve linear momentum and arise only from static deformations. Using continuum equations, we demonstrate the existence of the non-Hermitian skin effect in which the boundary hosts an extensive number of localized modes. Furthermore, lattice models reveal non-Hermitian topological transitions mediated by exceptional rings driven by the activity level of individual bonds.

Topological Waves in Fluids with Odd Viscosity.

Fluids in which both time reversal and parity are broken can display a dissipationless viscosity that is odd under each of these symmetries. Here, we show how this odd viscosity has a dramatic effect on topological sound waves in fluids, including the number and spatial profile of topological edge modes. Odd viscosity provides a short-distance cutoff that allows us to define a bulk topological invariant on a compact momentum space. As the sign of odd viscosity changes, a topological phase transition occurs without closing the bulk gap. Instead, at the transition point, the topological invariant becomes ill defined because momentum space cannot be compactified. This mechanism is unique to continuum models and can describe fluids ranging from electronic to chiral active systems.

Topological Protection Can Arise from Thermal Fluctuations and Interactions.

Topological quantum and classical materials can exhibit robust properties that are protected against disorder, for example, for noninteracting particles and linear waves. Here, we demonstrate how to construct topologically protected states that arise from the combination of strong interactions and thermal fluctuations inherent to soft materials or miniaturized mechanical structures. Specifically, we consider fluctuating lines under tension (e.g., polymer or vortex lines), subject to a class of spatially modulated substrate potentials. At equilibrium, the lines acquire a collective tilt proportional to an integer topological invariant called the Chern number. This quantized tilt is robust against substrate disorder, as verified by classical Langevin dynamics simulations. This robustness arises because excitations in this system of thermally fluctuating lines are gapped by virtue of interline interactions. We establish the topological underpinning of this pattern via a mapping that we develop between the interacting-lines system and a hitherto unexplored generalization of Thouless pumping to imaginary time. Our work points to a new class of classical topological phenomena in which the topological signature manifests itself in a structural property observed at finite temperature rather than a transport measurement.

A mechanism for anomalous transport in chiral active liquids.

Chiral active fluids are known to have anomalous transport properties such as the so-called odd viscosity. In this paper, we provide a microscopic mechanism for how such anomalous transport coefficients can emerge. We construct an Irving-Kirkwood-type stress tensor for chiral liquids and express the transport coefficients in terms of orientation-averaged intermolecular forces and distortions of the pair correlation function induced by a flow field. We then show how anomalous transport properties can be expected naturally due to the presence of a transverse component in the orientation-averaged intermolecular forces and anomalous distortion modes of the pair correlation function between chiral active particles. We anticipate that our work can provide a microscopic framework to explain the transport properties of nonequilibrium chiral systems.