Machine learning topological defects in confluent tissues.

Active nematics is an emerging paradigm for characterizing biological systems. One aspect of particularly intense focus is the role active nematic defects play in these systems, as they have been found to mediate a growing number of biological processes. Accurately detecting and classifying these defects in biological systems is, therefore, of vital importance to improving our understanding of such processes. While robust methods for defect detection exist for systems of elongated constituents, other systems, such as epithelial layers, are not well suited to such methods. Here, we address this problem by developing a convolutional neural network to detect and classify nematic defects in confluent cell layers. Crucially, our method is readily implementable on experimental images of cell layers and is specifically designed to be suitable for cells that are not rod shaped, which we demonstrate by detecting defects on experimental data using the trained model. We show that our machine learning model outperforms current defect detection techniques and that this manifests itself in our method as requiring less data to accurately capture defect properties. This could drastically improve the accuracy of experimental data interpretation while also reducing costs, advancing the study of nematic defects in biological systems.

Solving the stochastic dynamics of population growth.

Population growth is a fundamental process in ecology and evolution. The population size dynamics during growth are often described by deterministic equations derived from kinetic models. Here, we simulate several population growth models and compare the size averaged over many stochastic realizations with the deterministic predictions. We show that these deterministic equations are generically bad predictors of the average stochastic population dynamics. Specifically, deterministic predictions overestimate the simulated population sizes, especially those of populations starting with a small number of individuals. Describing population growth as a stochastic birth process, we prove that the discrepancy between deterministic predictions and simulated data is due to unclosed-moment dynamics. In other words, the deterministic approach does not consider the variability of birth times, which is particularly important with small population sizes. We show that some moment-closure approximations describe the growth dynamics better than the deterministic prediction. However, they do not reduce the error satisfactorily and only apply to some population growth models. We explicitly solve the stochastic growth dynamics, and our solution applies to any population growth model. We show that our solution exactly quantifies the dynamics of a community composed of different strains and correctly predicts the fixation probability of a strain in a serial dilution experiment. Our work sets the foundations for a more faithful modeling of community and population dynamics. It will allow the development of new tools for a more accurate analysis of experimental and empirical results, including the inference of important growth parameters.

Irreversibility across a Nonreciprocal PT-Symmetry-Breaking Phase Transition.

Nonreciprocal interactions are commonplace in continuum-level descriptions of both biological and synthetic active matter, yet studies addressing their implications for time reversibility have so far been limited to microscopic models. Here, we derive a general expression for the average rate of informational entropy production in the most generic mixture of conserved phase fields with nonreciprocal couplings and additive conservative noise. For the particular case of a binary system with Cahn-Hilliard dynamics augmented by nonreciprocal cross-diffusion terms, we observe a nontrivial scaling of the entropy production rate across a parity-time symmetry breaking phase transition. We derive a closed-form analytic expression in the weak-noise regime for the entropy production rate due to the emergence of a macroscopic dynamic phase, showing it can be written in terms of the global polar order parameter, a measure of parity-time symmetry breaking.

Intermittent attractive interactions lead to microphase separation in nonmotile active matter.

Nonmotile active matter exhibits a wide range of nonequilibrium collective phenomena yet examples are crucially lacking in the literature. We present a microscopic model inspired by the bacteria Neisseria meningitidis in which diffusive agents feel intermittent attractive forces. Through a formal coarse-graining procedure, we show that this truly scalar model of active matter exhibits the time-reversal-symmetry breaking terms defining the Active Model B+ class. In particular, we confirm the presence of microphase separation by solving the kinetic equations numerically. We show that the switching rate controlling the interactions provides a regulation mechanism tuning the typical cluster size, e.g., in populations of bacteria interacting via type IV pili.

Polar Fluctuations Lead to Extensile Nematic Behavior in Confluent Tissues.

How can a collection of motile cells, each generating contractile nematic stresses in isolation, become an extensile nematic at the tissue level? Understanding this seemingly contradictory experimental observation, which occurs irrespective of whether the tissue is in the liquid or solid states, is not only crucial to our understanding of diverse biological processes, but is also of fundamental interest to soft matter and many-body physics. Here, we resolve this cellular to tissue level disconnect in the small fluctuation regime by using analytical theories based on hydrodynamic descriptions of confluent tissues, in both liquid and solid states. Specifically, we show that a collection of microscopic constituents with no inherently nematic extensile forces can exhibit active extensile nematic behavior when subject to polar fluctuating forces. We further support our findings by performing cell level simulations of minimal models of confluent tissues.

Adhesion as a trigger of droplet polarization in flowing emulsions.

Tissues are subjected to large external forces and undergo global deformations during morphogenesis. We use synthetic analogues of tissues to study the impact of cell-cell adhesion on the response of cohesive cellular assemblies under such stresses. In particular, we use biomimetic emulsions in which the droplets are functionalized in order to exhibit specific droplet-droplet adhesion. We flow these emulsions in microfluidic constrictions and study their response to this forced deformation via confocal microscopy. We find that the distributions of avalanche sizes are conserved between repulsive and adhesive droplets. However, adhesion locally impairs the rupture of droplet-droplet contacts, which in turn pulls on the rearranging droplets. As a result, adhesive droplets are a lot more deformed along the axis of elongation in the constriction. This finding could shed light on the origin of polarization processes during morphogenesis.

Depletion attraction impairs the plasticity of emulsions flowing in a constriction.

We study the elasto-plastic behavior of dense attractive emulsions under a mechanical perturbation. The attraction is introduced through non-specific depletion interactions between the droplets and is controlled by changing the concentration of surfactant micelles in the continuous phase. We find that such attractive forces are not sufficient to induce any measurable modification on the scalings between the local packing fraction and the deformation of the droplets. However, when the emulsions are flowed through 2D microfluidic constrictions, we uncover a measurable effect of attraction on their elasto-plastic response. Indeed, we measure higher levels of deformation inside the constriction for attractive droplets. In addition, we show that these measurements correlate with droplet rearrangements that are spatially delayed in the constriction for higher attraction forces.

Active acoustic switches using two-dimensional granular crystals.

We employ numerical simulations to study active transistor-like switches made from two-dimensional (2D) granular crystals containing two types of grains with the same size but different masses. We tune the mass contrast and arrangement of the grains to maximize the width of the frequency band gap in the device. The input signal is applied to a single grain on one side of the device, and the output signal is measured from another grain on the other side of the device. Changing the size of one or many grains tunes the pressure, which controls the vibrational response of the device. Switching between the on and off states is achieved using two mechanisms: (1) pressure-induced switching where the interparticle contact network is the same in the on and off states and (2) switching through contact breaking. In general, the performance of the acoustic switch, as captured by the gain ratio and switching time between the on and off states, is better for pressure-induced switching. We show that in these acoustic switches the gain ratio between the on and off states can be larger than 10^{4} and the switching time (multiplied by the driving frequency) is comparable to that obtained recently for sonic crystals and less than that for photonic transistor-like switches. Since the self-assembly of grains with different masses into 2D granular crystals is challenging, we describe simulations of circular grains with small circular knobs placed symmetrically around the perimeter mixed with circular grains without knobs. Using umbrella sampling techniques, we show that grains with six knobs most efficiently form the hexagonal crystals that yield the largest frequency band gap. Using the simulation results, we estimate the time required for vibration experiments to generate granular crystals of millimeter-sized steel beads with maximal band gaps.

Optimized Diffusion of Run-and-Tumble Particles in Crowded Environments.

We study the transport of self-propelled particles in dynamic complex environments. To obtain exact results, we introduce a model of run-and-tumble particles (RTPs) moving in discrete time on a d-dimensional cubic lattice in the presence of diffusing hard-core obstacles. We derive an explicit expression for the diffusivity of the RTP, which is exact in the limit of low density of fixed obstacles. To do so, we introduce a generalization of Kac's theorem on the mean return times of Markov processes, which we expect to be relevant for a large class of lattice gas problems. Our results show the diffusivity of RTPs to be nonmonotonic in the tumbling probability for low enough obstacle mobility. These results prove the potential for the optimization of the transport of RTPs in crowded and disordered environments with applications to motile artificial and biological systems.

Response of jammed packings to thermal fluctuations.

We focus on the response of mechanically stable (MS) packings of frictionless, bidisperse disks to thermal fluctuations, with the aim of quantifying how nonlinearities affect system properties at finite temperature. In contrast, numerous prior studies characterized the structural and mechanical properties of MS packings of frictionless spherical particles at zero temperature. Packings of disks with purely repulsive contact interactions possess two main types of nonlinearities, one from the form of the interaction potential (e.g., either linear or Hertzian spring interactions) and one from the breaking (or forming) of interparticle contacts. To identify the temperature regime at which the contact-breaking nonlinearities begin to contribute, we first calculated the minimum temperatures T_{cb} required to break a single contact in the MS packing for both single- and multiple-eigenmode perturbations of the T=0 MS packing. We find that the temperature required to break a single contact for equal velocity-amplitude perturbations involving all eigenmodes approaches the minimum value obtained for a perturbation in the direction connecting disk pairs with the smallest overlap. We then studied deviations in the constant volume specific heat C[over ¯]_{V} and deviations of the average disk positions Δr from their T=0 values in the temperature regime T_{C[over ¯]_{V}}100 for linear spring interactions is independent of system size. This result emphasizes that contact-breaking nonlinearities are dominant over form nonlinearities in the low-temperature range T_{cb}

Local and global avalanches in a two-dimensional sheared granular medium.

We present the experimental and numerical studies of a two-dimensional sheared amorphous material composed of bidisperse photoelastic disks. We analyze the statistics of avalanches during shear including the local and global fluctuations in energy and changes in particle positions and orientations. We find scale-free distributions for these global and local avalanches denoted by power laws whose cutoffs vary with interparticle friction and packing fraction. Different exponents are found for these power laws depending on the quantity from which variations are extracted. An asymmetry in time of the avalanche shapes is evidenced along with the fact that avalanches are mainly triggered by the shear bands. A simple relation independent of the intensity is found between the number of local avalanches and the global avalanches they form. We also compare these experimental and numerical results for both local and global fluctuations to predictions from mean-field and depinning theories.

Protocol dependence of the jamming transition.

We propose a theoretical framework for predicting the protocol dependence of the jamming transition for frictionless spherical particles that interact via repulsive contact forces. We study isostatic jammed disk packings obtained via two protocols: isotropic compression and simple shear. We show that for frictionless systems, all jammed packings can be obtained via either protocol. However, the probability to obtain a particular jammed packing depends on the packing-generation protocol. We predict the average shear strain required to jam initially unjammed isotropically compressed packings from the density of jammed packings, shape of their basins of attraction, and path traversed in configuration space. We compare our predictions to simulations of shear strain-induced jamming and find quantitative agreement. We also show that the packing fraction range, over which shear strain-induced jamming occurs, tends to zero in the large system limit for frictionless packings with overdamped dynamics.

Hypocoordinated solids in particulate media.

We propose a "phase diagram" for particulate systems with purely repulsive contact forces, such as granular media and colloids. We characterize two classes of behavior as a function of the input kinetic energy per degree of freedom T_{0} and packing fraction deviation from jamming onset Δϕ=ϕ-ϕ_{J} using simulations of frictionless disks. Isocoordinated solids (ICS) exist above jamming; they possess an average contact number equal to the isostatic value z_{iso}. ICS display "strict" harmonic response, where the density of vibrational modes from the Fourier transform of the velocity autocorrelation function is a set of sharp peaks at eigenfrequencies ω_{k}{d} of the dynamical matrix. In contrast, hypocoordinated solids (HCS) occur above and below jamming and possess fluctuating networks of interparticle contacts but do not undergo cage-breaking particle rearrangements. The density of vibrational frequencies for the HCS is not a collection of sharp peaks at ω_{k}{d}, but it does possess a common form over a range of Δϕ and T_{0}.

Repulsive contact interactions make jammed particulate systems inherently nonharmonic.

Many jammed particulate systems, such as granular and colloidal materials, interact via repulsive contact forces. We find that these systems possess no harmonic regime in the large system limit (N→∞) for all compressions Δϕ studied, and at jamming onset Δϕ→0 for all N. We perform fixed energy simulations following perturbations with amplitude δ along eigendirections of the dynamical matrix. The fluctuations abruptly spread to all modes for δ≈δ(c) (where a single contact breaks) in contrast to linear and weakly nonlinear behavior. For δ > δ(c), all discrete modes disappear into a continuous frequency band. <δ(c)> scales with 1/N and Δϕ, which limits harmonic behavior to only overcompressed systems. The density of vibrational modes deviates strongly from that predicted from the dynamical matrix when the system enters the nonharmonic regime, which significantly affects its mechanical and transport properties.