Multicellular rosettes drive fluid-solid transition in epithelial tissues

Models for confluent biological tissues often describe the network formed by cells as a triple-junction network, similar to foams. However, higher order vertices or multicellular rosettes are prevalent in developmental and {\it in vitro} processes and have been recognized as crucial in many important aspects of morphogenesis, disease, and physiology. In this work, we study the influence of rosettes on the mechanics of a confluent tissue. We find that the existence of rosettes in a tissue can greatly influence its rigidity. Using a generalized vertex model and effective medium theory we find a fluid-to-solid transition driven by rosette density and intracellular tensions. This transition exhibits several hallmarks of a second-order phase transition such as a growing correlation length and a universal critical scaling in the vicinity a critical point. Further, we elucidate the nature of rigidity transitions in dense biological tissues and other cellular structures using a generalized Maxwell constraint counting approach. This answers a long-standing puzzle of the origin of solidity in these systems.Comment: 11 pages, 5 figures + 8 pages, 7 figures in Appendix. To be appear in PR

The Statistical Physics of Athermal Materials

At the core of equilibrium statistical mechanics lies the notion of statistical ensembles: a collection of microstates, each occurring with a given a priori probability that depends only on a few macroscopic parameters such as temperature, pressure, volume, and energy. In this review article, we discuss recent advances in establishing statistical ensembles for athermal materials. The broad class of granular and particulate materials is immune from the effects of thermal fluctuations because the constituents are macroscopic. In addition, interactions between grains are frictional and dissipative, which invalidates the fundamental postulates of equilibrium statistical mechanics. However, granular materials exhibit distributions of microscopic quantities that are reproducible and often depend on only a few macroscopic parameters. We explore the history of statistical ensemble ideas in the context of granular materials, clarify the nature of such ensembles and their foundational principles, highlight advances in testing key ideas, and discuss applications of ensembles to analyze the collective behavior of granular materials

Fluctuations in Shear-Jammed States: A Statistical Ensemble Approach

Granular matter exists out of thermal equilibrium, i.e. it is athermal. While conventional equilibrium statistical mechanics is not useful for characterizing granular materials, the idea of constructing a statistical ensemble analogous to its equilibrium counterpart to describe static granular matter was proposed by Edwards and Oakshott more than two decades ago. Recent years have seen several implementations of this idea. One of these is the stress ensemble, which is based on properties of the force moment tensor, and applies to frictional and frictionless grains. We demonstrate the full utility of this statistical framework in shear jammed (SJ) experimental states [1,2], a special class of granular solids created by pure shear, which is a strictly non-equilbrium protocol for creating solids. We demonstrate that the stress ensemble provides an excellent quantitative description of fluctuations in experimental SJ states. We show that the stress fluctuations are controlled by a single tensorial quantity: the angoricity of the system, which is a direct analog of the thermodynamic temperature. SJ states exhibit significant correlations in local stresses and are thus inherently different from density-driven, isotropically jammed (IJ) states.Comment: 6 pages, 4 figure

Motility-driven glass and jamming transitions in biological tissues

Cell motion inside dense tissues governs many biological processes, including embryonic development and cancer metastasis, and recent experiments suggest that these tissues exhibit collective glassy behavior. To make quantitative predictions about glass transitions in tissues, we study a self-propelled Voronoi (SPV) model that simultaneously captures polarized cell motility and multi-body cell-cell interactions in a confluent tissue, where there are no gaps between cells. We demonstrate that the model exhibits a jamming transition from a solid-like state to a fluid-like state that is controlled by three parameters: the single-cell motile speed, the persistence time of single-cell tracks, and a target shape index that characterizes the competition between cell-cell adhesion and cortical tension. In contrast to traditional particulate glasses, we are able to identify an experimentally accessible structural order parameter that specifies the entire jamming surface as a function of model parameters. We demonstrate that a continuum Soft Glassy Rheology model precisely captures this transition in the limit of small persistence times, and explain how it fails in the limit of large persistence times. These results provide a framework for understanding the collective solid-to-liquid transitions that have been observed in embryonic development and cancer progression, which may be associated with Epithelial-to-Mesenchymal transition in these tissues.Comment: accepted for publication in Physical Review X, 201

Controlled neighbor exchanges drive glassy behavior, intermittency and cell streaming in epithelial tissues

Cell neighbor exchanges are integral to tissue rearrangements in biology, including development and repair. Often these processes occur via topological T1 transitions analogous to those observed in foams, grains and colloids. However, in contrast to in non-living materials the T1 transitions in biological tissues are rate-limited and cannot occur instantaneously due to the finite time required to remodel complex structures at cell-cell junctions. Here we study how this rate-limiting process affects the mechanics and collective behavior of cells in a tissue by introducing this important biological constraint in a theoretical vertex-based model as an intrinsic single-cell property. We report in the absence of this time constraint, the tissue undergoes a motility-driven glass transition characterized by a sharp increase in the intermittency of cell-cell rearrangements. Remarkably, this glass transition disappears as T1 transitions are temporally limited. As a unique consequence of limited rearrangements, we also find that the tissue develops spatially correlated streams of fast and slow cells, in which the fast cells organize into stream-like patterns with leader-follower interactions, and maintain optimally stable cell-cell contacts. The predictions of this work is compared with existing in-vivo experiments in Drosophila pupal development

Why Do Granular Materials Stiffen with Shear Rate? : Test of Novel Stress-Based Statistics

Peer reviewedPublisher PD

Shear-induced rigidity of frictional particles: Analysis of emergent order in stress space

Solids are distinguished from fluids by their ability to resist shear. In traditional solids, the resistance to shear is associated with the emergence of broken translational symmetry as exhibited by a non-uniform density pattern, which results from either minimizing the energy cost or maximizing the entropy or both. In this work, we focus on a class of systems, where this paradigm is challenged. We show that shear-driven jamming in dry granular materials is a collective process controlled solely by the constraints of mechanical equilibrium. We argue that these constraints lead to a broken translational symmetry in a dual space that encodes the statistics of contact forces and the topology of the contact network. The shear-jamming transition is marked by the appearance of this broken symmetry. We extend our earlier work, by comparing and contrasting real space measures of rheology with those obtained from the dual space. We investigate the structure and behavior of the dual space as the system evolves through the rigidity transition in two different shear protocols. We analyze the robustness of the shear-jamming scenario with respect to protocol and packing fraction, and demonstrate that it is possible to define a protocol-independent order parameter in this dual space, which signals the onset of rigidity.Comment: 14 pages, 17 figure