Spontaneous crumpling of active spherical shells.

The existence of a crumpled phase for self-avoiding elastic surfaces was postulated more than three decades ago using simple Flory-like scaling arguments. Despite much effort, its stability in a microscopic environment has been the subject of much debate. In this paper we show how a crumpled phase develops reliably and consistently upon subjecting a thin spherical shell to active fluctuations. We find a master curve describing how the relative volume of a shell changes with the strength of the active forces, that applies for every shell independent of size and elastic constants. Furthermore, we extract a general expression for the onset active force beyond which a shell begins to crumple. Finally, we calculate how the size exponent varies along the crumpling curve.

The crumpling transition of active tethered membranes.

We perform numerical simulations of active ideal and self-avoiding tethered membranes. Passive ideal membranes with bending interactions are known to exhibit a continuous crumpling transition between a low temperature flat phase and a high temperature crumpled phase. Conversely, self-avoiding membranes remain in an extended (flat) phase for all temperatures even in the absence of a bending energy. We find that the introduction of active fluctuations into the system produces a phase behavior that is overall consistent with that observed for passive membranes. The phases and the nature of the transition for ideal membranes is unchanged and active fluctuations can be remarkably accounted for by a simple rescaling of the temperature. For the self-avoiding membrane, we find that the extended phase is preserved even in the presence of very large active fluctuations.

Rectification of confined soft vesicles containing active particles.

One of the most promising features of active systems is that they can extract energy from their environment and convert it to mechanical work. Self propelled particles enable rectification when in contact with rigid boundaries. They can rectify their own motion when confined in asymmetric channels and that of microgears. In this paper, we study the shape fluctuations of two dimensional flexible vesicles containing active Brownian particles. We show how these fluctuations not only are capable of easily squeezing a vesicle through narrow openings, but are also responsible for its rectification when placed within asymmetric confining channels (ratchetaxis). We detail the conditions under which this process can be optimized, and sort out the complex interplay between elastic and active forces responsible for the directed motion of the vesicle across these channels.

Loops versus lines and the compression stiffening of cells.

Both animal and plant tissue exhibit a nonlinear rheological phenomenon known as compression stiffening, or an increase in moduli with increasing uniaxial compressive strain. Does such a phenomenon exist in single cells, which are the building blocks of tissues? One expects an individual cell to compression soften since the semiflexible biopolymer-based cytoskeletal network maintains the mechanical integrity of the cell and in vitro semiflexible biopolymer networks typically compression soften. To the contrary, we find that mouse embryonic fibroblasts (mEFs) compression stiffen under uniaxial compression via atomic force microscopy studies. To understand this finding, we uncover several potential mechanisms for compression stiffening. First, we study a single semiflexible polymer loop modeling the actomyosin cortex enclosing a viscous medium modeled as an incompressible fluid. Second, we study a two-dimensional semiflexible polymer/fiber network interspersed with area-conserving loops, which are a proxy for vesicles and fluid-based organelles. Third, we study two-dimensional fiber networks with angular-constraining crosslinks, i.e. semiflexible loops on the mesh scale. In the latter two cases, the loops act as geometric constraints on the fiber network to help stiffen it via increased angular interactions. We find that the single semiflexible polymer loop model agrees well with the experimental cell compression stiffening finding until approximately 35% compressive strain after which bulk fiber network effects may contribute. We also find for the fiber network with area-conserving loops model that the stress-strain curves are sensitive to the packing fraction and size distribution of the area-conserving loops, thereby creating a mechanical fingerprint across different cell types. Finally, we make comparisons between this model and experiments on fibrin networks interlaced with beads as well as discuss implications for single cell compression stiffening at the tissue scale.

Effective forces between active polymers.

The characterization of the interactions between two fully flexible self-avoiding polymers is one of the classic and most important problems in polymer physics. In this paper we measure these interactions in the presence of active fluctuations. We introduce activity into the problem using two of the most popular models in this field, one where activity is effectively embedded into the monomers' dynamics, and the other where passive polymers fluctuate in an explicit bath of active particles. We establish the conditions under which the interaction between active polymers can be mapped into the classical passive problem. We observe that the active bath can drive the development of strong attractive interactions between the polymers and that, upon enforcing a significant degree of overlap, they come together to form a single double-stranded unit. A phase diagram tracing this change in conformational behavior is also reported.

Rigidity transitions in zero-temperature polygons.

We study geometrical clues of a rigidity transition due to the emergence of a system-spanning state of self-stress in underconstrained systems of individual polygons and spring networks constructed from such polygons. When a polygon with harmonic bond edges and an area spring constraint is subject to an expansive strain, we observe that convexity of the polygon is a necessary condition for such a self-stress. We prove that the cyclic configuration of the polygon is a sufficient condition for the self-stress. This correspondence of geometry and rigidity is akin to the straightening of a one dimensional chain of springs to rigidify it. We predict the onset of the rigidity transition and estimate the transition strain using purely geometrical methods. These findings help determine the rigidity of an area-preserving polygon just by knowing its geometry. Since two-dimensional spring networks can be considered as a network of polygons, we look for similar geometric features in underconstrained spring networks under isotropic expansive strain. We observe that all polygons attain convexity at the rigidity transition such that the fraction of convex, but not cyclic, polygons predicts the onset of the rigidity transition. Acyclic polygons in the network correlate with larger tensions, forming effective force chains.