J. M. Schwarz

A density-independent rigidity transition in biological tissues

Cells moving in a tissue undergo a rigidity transition resembling that of active particles jamming at a critical density—but the tissue density stays constant. A new type of rigidity transition implicates the physical properties of the cells.

Energy barriers govern glassy dynamics in tissues

Recent observations demonstrate that confluent tissues exhibit features of glassy dynamics, such as caging behavior and dynamical heterogeneities, although it has remained unclear how single-cell properties control this behavior. Here we develop numerical and theoretical models to calculate energy barriers to cell rearrangements, which help govern cell migration in cell monolayers. In contrast to work on sheared foams, we find that energy barrier heights are exponentially distributed and depend systematically on the cells number of neighbors. Based on these results, we predict glassy two-time correlation functions for cell motion, with a timescale that increases rapidly as cell activity decreases. These correlation functions are used to construct simple random walks that reproduce the caging behavior observed for cell trajectories in experiments. This work provides a theoretical framework for predicting collective motion of cells in wound-healing, embryogenesis and cancer tumorogenesis.

Redundancy and cooperativity in the mechanics of compositely crosslinked filamentous networks.

The cytoskeleton of living cells contains many types of crosslinkers. Some crosslinkers allow energy-free rotations between filaments and others do not. The mechanical interplay between these different crosslinkers is an open issue in cytoskeletal mechanics. Therefore, we develop a theoretical framework based on rigidity percolation to study a generic filamentous system containing both stretching and bond-bending forces to address this issue. The framework involves both analytical calculations via effective medium theory and numerical simulations on a percolating triangular lattice with very good agreement between both. We find that the introduction of angle-constraining crosslinkers to a semiflexible filamentous network with freely rotating crosslinks can cooperatively lower the onset of rigidity to the connectivity percolation threshold—a result argued for years but never before obtained via effective medium theory. This allows the system to ultimately attain rigidity at the lowest concentration of material possible. We further demonstrate that introducing angle-constraining crosslinks results in mechanical behaviour similar to just freely rotating crosslinked semflexible filaments, indicating redundancy and universality. Our results also impact upon collagen and fibrin networks in biological and bio-engineered tissues.

Branching, capping, and severing in dynamic actin structures.

Branched actin networks at the leading edge of a crawling cell evolve via protein-regulated processes such as polymerization, depolymerization, capping, branching, and severing. A formulation of these processes is presented and analyzed to study steady-state network morphology. In bulk, we identify several scaling regimes in severing and branching protein concentrations and find that the coupling between severing and branching is optimally exploited for conditions in vivo. Near the leading edge, we find qualitative agreement with the in vivo morphology.

Driven Depinning of Strongly Disordered Media and Anisotropic Mean-Field Limits

Extended systems driven through strong disorder are modeled generically using coarse-grained degrees of freedom that interact elastically in the directions parallel to the drive and slip along at least one of the directions transverse to the motion. In the limit of infinite-range elastic and viscous coupling this model has a tricritical point separating a region where the depinning is continuous, in the universality class of elastic depinning, from a region where depinning is hysteretic. Many of the collective transport models discussed in the literature are special cases of the generic model.

Rigid Cluster Decomposition Reveals Criticality in Frictional Jamming

We study the nature of the frictional jamming transition within the framework of rigidity percolation theory. Slowly sheared frictional packings are decomposed into rigid clusters and floppy regions with a generalization of the pebble game including frictional contacts. Our method suggests a second-order transition controlled by the emergence of a system-spanning rigid cluster accompanied by a critical cluster size distribution. Rigid clusters also correlate with common measures of rigidity. We contrast this result with frictionless jamming, where the rigid cluster size distribution is noncritical.

On the Modeling of Endocytosis in Yeast

The cell membrane deforms during endocytosis to surround extracellular material and draw it into the cell. Results of experiments on endocytosis in yeast show general agreement that 1) actin polymerizes into a network of filaments exerting active forces on the membrane to deform it, and 2) the large-scale membrane deformation is tubular in shape. In contrast, there are three competing proposals for precisely how the actin filament network organizes itself to drive the deformation. We use variational approaches and numerical simulations to address this competition by analyzing a meso-scale model of actin-mediated endocytosis in yeast. The meso-scale model breaks up the invagination process into three stages: 1) initiation, where clathrin interacts with the membrane via adaptor proteins; 2) elongation, where the membrane is then further deformed by polymerizing actin filaments; and 3) pinch-off. Our results suggest that the pinch-off mechanism may be assisted by a pearling-like instability. We rule out two of the three competing proposals for the organization of the actin filament network during the elongation stage. These two proposals could be important in the pinch-off stage, however, where additional actin polymerization helps break off the vesicle. Implications and comparisons with earlier modeling of endocytosis in yeast are discussed.

Jamming graphs: A local approach to global mechanical rigidity

We revisit the concept of minimal rigidity as applied to frictionless, repulsive soft sphere packings in two dimensions with the introduction of the jamming graph. Minimal rigidity is a purely combinatorial property encoded via Lamans theorem in two dimensions. It constrains the global, average coordination number of the graph, for example. However, minimal rigidity does not address the geometry of local mechanical stability. The jamming graph contains both properties of global mechanical stability at the onset of jamming and local mechanical stability. We demonstrate how jamming graphs can be constructed using local moves via the Henneberg construction such that these graphs fall under the jurisdiction of correlated percolation. We then probe how jamming graphs destabilize, or become unjammed, by deleting a bond and computing the resulting rigid cluster distribution. We also study how the system restabilizes with the addition of new contacts and how a jamming graph with extra (redundant) contacts destabilizes. The latter endeavor allows us to probe a disk packing in the rigid phase and uncover a potentially new diverging length scale associated with the random deletion of contacts as compared to the study of cut-out (or frozen-in) subsystems.

Mean-field theory of collective transport with phase slips

The driven transport of plastic systems in various disordered backgrounds is studied within mean field theory. Plasticity is modeled using nonconvex interparticle potentials that allow for phase slips. This theory most naturally describes sliding charge density waves; other applications include flow of colloidal particles or driven magnetic flux vortices in disordered backgrounds. The phase diagrams exhibit generic phases and phase boundaries, though the shapes of the phase boundaries depend on the shape of the disorder potential. The phases are distinguished by their velocity and coherence: the moving phase generically has finite coherence, while pinned states can be coherent or incoherent. The coherent and incoherent static phases can coexist in parameter space, in contrast with previous results for exactly sinusoidal pinning potentials. Transitions between the moving and static states can also be hysteretic. The depinning transition from the static to sliding states can be determined analytically, while the repinning transition from the moving to the pinned phases is computed by direct simulation.

Shape-Shifting Droplet Networks.

We consider a three-dimensional network of aqueous droplets joined by single lipid bilayers to form a cohesive, tissuelike material. The droplets in these networks can be programed to have distinct osmolarities so that osmotic gradients generate internal stresses via local fluid flows to cause the network to change shape. We discover, using molecular dynamics simulations, a reversible folding-unfolding process by adding an osmotic interaction with the surrounding environment which necessarily evolves dynamically as the shape of the network changes. This discovery is the next important step towards osmotic robotics in this system. We also explore analytically and numerically how the networks become faceted via buckling and how quasi-one-dimensional networks become three dimensional.