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We investigate the emergence of orientational order among +1/2 disclinations in active nematic liquid crystals. Using a combination of theoretical and experimental methods, we show that +1/2 disclinations have short-range antiferromagnetic alignment, as a consequence of the elastic torques originating from their polar structure. The presence of intermediate -1/2 disclinations, however, turns this interaction from anti-aligning to aligning at scales that are smaller than the typical distance between like-sign defects. No long-range orientational order is observed. Strikingly, these effects are insensitive to material properties and qualitatively similar to what is found for defects in passive nematic liquid crystals., 6 pages, 4 figures

Topological defects are one of the most conspicuous features of liquid crystals. In two dimensional nematics, they have been shown to behave effectively as particles with both, charge and orientation, which dictate their interactions. Here, we study "twisted" defects that have a radially dependent orientation. We find that twist can be partially relaxed through the creation and annihilation of defect pairs. By solving the equations for defect motion and calculating the forces on defects, we identify four distinct elements that govern the relative relaxational motion of interacting topological defects, namely attraction, repulsion, co-rotation and co-translation. The interaction of these effects can lead to intricate defect trajectories, which can be controlled by setting relevant timescales., 8 pages, 6 figures

We investigate the emergence of global alignment in colonies of dividing rod-shaped cells under confinement. Using molecular dynamics simulations and continuous modeling, we demonstrate that geometrical anisotropies in the confining environment give rise to imbalance in the normal stresses, which, in turn, drives a collective rearrangement of the cells. This behavior crucially relies on the colony's solid-like mechanical response at short time scales and can be recovered within the framework of active hydrodynamics upon modeling bacterial colonies as growing viscoelastic gels characterized by Maxwell-like stress relaxation., Comment: 10 pages, 8 figures

We investigate the effects of extrinsic curvature on the turbulent behavior of a 2D active nematic confined to the surface of a cylinder. The surface of a cylinder has no intrinsic curvatrue and only extrinsic curvature. A nematic field reacts to the extrinsic curvature by trying to align with the lowest principle curvature, in this case parallel to the long axis of the cylinder. When nematics are sufficiently active, there is a proliferation of defects arising from a bend or splay instability depending on the nature of the active stress. The extrinsic curvature of the cylinder beaks the rotational symmetry of this process, implying that defects are created parallel or perpendicular to the cylinder depending on whether the active nematic is contractile or extensile., Comment: 6 pages, 7 Figures

The transition from monolayers to multilayered structures in bacterial colonies is a fundamental step in biofilm development. Observed across different morphotypes and species, this transition is triggered within freely growing bacterial microcolonies comprising a few hundred cells. Using a combination of numerical simulations and analytical modeling, here we demonstrate that this transition originates from the competition between growth-induced in-plane active stresses and vertical restoring forces, due to the cell-substrate interactions. Using a simple chainlike colony of laterally confined cells, we show that the transition sets when individual cells become unstable to rotations, thus it is localized and mechanically deterministic. Asynchronous cell division renders the process stochastic, so that all the critical parameters that control the onset of the transition are continuously distributed random variables. Here we demonstrate that the occurrence of the first division in the colony can be approximated as a Poisson process in the limit of large cells numbers. This allows us to approximately calculate the probability distribution function of the position and time associated with the first extrusion. The rate of such a Poisson process can be identified as the order parameter of the transition, thus highlighting its mixed deterministic-stochastic nature., 5 pages, 4 figures

We investigate the turbulent dynamics of a two-dimensional active nematic liquid crystal con- strained on a curved surface. Using a combination of hydrodynamic and particle-based simulations, we demonstrate that the fundamental structural features of the fluid, such as the topological charge density, the defect number density, the nematic order parameter and defect creation and annihilation rates, are simple linear functions of the substrate Gaussian curvature, which then acts as a control parameter for the chaotic flow. Our theoretical predictions are then compared with experiments on microtubule-kinesin suspensions confined on toroidal active droplets, finding excellent qualitative agreement., Comment: 6 pages, 4 figures

For bacterial cell division, treadmilling filaments of FtsZ organize into a ring-like structure at the center of the cell. What governs the architecture and stability of this dynamic Z-ring is currently unknown, but FtsZ-associated proteins have been suggested to play an important role. Here, we used anin vitroreconstitution approach combined with fluorescence microscopy to study the influence of the well-conserved protein ZapA on the organization and dynamics of FtsZ filaments recruited to a supported membrane. We found that ZapA increases the spatial order and stabilizes the steady-state architecture of the FtsZ filament network in a highly cooperative manner. Despite its strong influence on their large-scale organization, ZapA binds only transiently to FtsZ filaments and has no effect on their treadmilling velocity. Together, our data explains how FtsZ-associated proteins can contribute to the precision and stability of the Z-ring without compromising treadmilling dynamics.

We investigate the effect of an anisotropic substrate on the turbulent dynamics of a simulated two dimensional active nematic. This is introduced as an anisotropic friction and an effective anisotropic viscosity, with the orientation of the anisotropy being defined by the substrate. In this system we observe the emergence of global nematic order of topological defects that is controlled by the degree of anisotropy in the viscosity and the magnitude of the active stress. No global defect alignment is seen in passive liquid crystals with anisotropic viscosity or friction confirming that ordering is driven by the active stress. We then closely examine the active flow generated by a single defect to show that the kinetic energy of the flow is orientation dependent, resulting in a torque on the defect to align them with the anisotropy in the substrate., Comment: 5 pages, 4 figures

Teams of cooperating sperm have been found across several vertebrate and invertebrate species, ranging from sperm pairs to massive aggregates containing hundreds of cells. Although the biochemical mechanisms involved in the aggregation process are still unclear, it was found that aggregation can enhance the mobility of the cells, thus offering an advantage during fertilization. Here, we report a thorough computational investigation on the role of cellular geometry in the performance of sperm aggregates. The sperm head is modelled as a persistent random walker characterized by a non-trivial three-dimensional shape and equipped with an adhesive region, where cell-cell binding occurs. By considering both a simple parametric head shape and a computer reconstruction of a real head shape based on morphometric data, we demonstrate that the geometry of the head and the structure of the adhesive region crucially affects both the stability and mobility of the aggregates. Our analysis further suggests that the apical hook commonly found in the sperm of muroid rodents, might serve to shield portions of the adhesive region and promote efficient alignment of the velocities of the interacting cells., 6 pages, 7 figures

Nematic order on curved surfaces is often disrupted by the presence of topological defects, which are singular regions in which the orientational order is undefined. In the presence of force-generating active materials, these defects are able to migrate through space like swimming microorganisms. We use toroidal surfaces to show that despite their highly chaotic and non-equilibrium dynamics, pairs of defects unbind and segregate in regions of opposite Gaussian curvature. Using numerical simulations, we find that the degree of defect unbinding can be controlled by tuning the system activity, and even suppressed in strongly active systems. Furthermore, by using the defects as active microrheological tracers and quantitatively comparing our experimental and theoretical results, we are able to determine material properties of the active nematic. Our results illustrate how topology and geometry can be used to control the behaviour of active materials, and introduce a new avenue for the quantitative mechanical characterization of active fluids. Topological defects in a turbulent active nematic on a toroidal surface are shown to segregate in regions of opposite curvature. Simulations suggest that this behaviour may be controlled — or even suppressed — by tuning the level of activity.

Swarming is a conspicuous behavioural trait observed in bird flocks, fish shoals, insect swarms and mammal herds. It is thought to improve collective awareness and offer protection from predators. Many current models involve the hypothesis that information coordinating motion is exchanged between neighbors. We argue that such local interactions alone are insufficient to explain the organization of large flocks of birds and that the mechanism for the exchange of long-ranged information necessary to control their density remains unknown. We show that large flocks self-organize to the maximum density at which a typical individual is still just able to see out of the flock in many directions. Such flocks are marginally opaque - an external observer can also just still see a substantial fraction of sky through the flock. Although seemingly intuitive we show that this need not be the case; flocks could easily be highly diffuse or entirely opaque. The emergence of marginal opacity strongly constrains how individuals interact with each other within large swarms. It also provides a mechanism for global interactions: An individual can respond to the projection of the flock that it sees. This provides for faster information transfer and hence rapid flock dynamics, another advantage over local models. From a behavioural perspective it optimizes the information available to each bird while maintaining the protection of a dense, coherent flock., PNAS early edition published online at http://www.pnas.org/cgi/doi/10.1073/pnas.1402202111

Bacterial colonies are abundant on living and nonliving surfaces and are known to mediate a broad range of processes in ecology, medicine, and industry. Although extensively researched, from single cells to demographic scales, a comprehensive biomechanical picture, highlighting the cell-to-colony dynamics, is still lacking. Here, using molecular dynamics simulations and continuous modeling, we investigate the geometrical and mechanical properties of a bacterial colony growing on a substrate with a free boundary and demonstrate that such an expanding colony self-organizes into a “mosaic” of microdomains consisting of highly aligned cells. The emergence of microdomains is mediated by two competing forces: the steric forces between neighboring cells, which favor cell alignment, and the extensile stresses due to cell growth that tend to reduce the local orientational order and thereby distort the system. This interplay results in an exponential distribution of the domain areas and sets a characteristic length scale proportional to the square root of the ratio between the system orientational stiffness and the magnitude of the extensile active stress. Our theoretical predictions are finally compared with experiments with freely growing E. coli microcolonies, finding quantitative agreement., Physical Review X, 8 (3), ISSN:2160-3308

Large collections of autonomously moving agents, such as animals or micro-organisms, are able to 'flock' coherently in space even in the absence of a central control mechanism. While the direction of the flock resulting from this critical behavior is random, this can be controlled by a small subset of informed individuals acting as leaders of the group. In this article we use the Vicsek model to investigate how flocks respond to leadership and make decisions. Using a combination of numerical simulations and continuous modeling we demonstrate that flocks display a linear response to leadership that can be cast in the framework of the fluctuation-dissipation theorem, identifying an 'effective temperature' reflecting how promptly the flock reacts to the initiative of the leaders. The linear response to leadership also holds in the presence of two groups of informed individuals with competing interests, indicating that the flock's behavioral decision is determined by both the number of leaders and their degree of influence., Comment: 8 pages (incl. supplementary information), 8 figures, Supplementary movies can be found at http://wwwhome.lorentz.leidenuniv.nl/~giomi/sup_mat/20151108/