Density regulation in strictly metric-free swarms

There is now experimental evidence that nearest-neighbour interactions in flocks of birds are metric free, i.e. they have no characteristic interaction length scale. However, models that involve interactions between neighbours that are assigned topologically are naturally invariant under spatial expansion, supporting a continuous reduction in density towards zero, unless additional cohesive interactions are introduced or the density is artificially controlled, e.g. via a finite system size. We propose a solution that involves a metric-free motional bias on those individuals that are topologically identified to be on an edge of the swarm. This model has only two primary control parameters, one controlling the relative strength of stochastic noise to the degree of co-alignment and another controlling the degree of the motional bias for those on the edge, relative to the tendency to co-align. We find a novel power-law scaling of the real-space density with the number of individuals N as well as a familiar order-to-disorder transition

Geometry and mechanics of microdomains in growing bacterial colonies

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.Comment: 10 pages, 7 figure

The Role of Projection in the Control of Bird Flocks

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.Comment: PNAS early edition published online at http://www.pnas.org/cgi/doi/10.1073/pnas.140220211

Emergent behavioural phenotypes of swarming models revealed by mimicking a frustrated anti-ferromagnet

Self-propelled particle (SPP) models are often compared with animal swarms. However, the collective animal behaviour observed in experiments often leaves considerable unconstrained freedom in the structure of a proposed model. Essentially, multiple models can describe the observed behaviour of animal swarms in simple environments. To tackle this degeneracy, we study swarms of SPPs in non-trivial environments as a new approach to distinguish between candidate models. We restrict swarms of SPPs to circular (periodic) channels where they polarize in one of two directions (like spins) and permit information to pass through windows between neighbouring channels. Co-alignment between particles then couples the channels (anti-ferromagnetically) so that they tend to counter-rotate. We study channels arranged to mimic a geometrically frustrated anti-ferromagnet and show how the effects of this frustration allow us to better distinguish between SPP models. Similar experiments could therefore improve our understanding of collective motion in animals. Finally, we discuss how the spin analogy can be exploited to construct universal logic gates, and therefore swarming systems that can function as Turing machines

Confinement-induced self-organization in growing bacterial colonies

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 figure

Mono-to-multilayer transition in growing bacterial colonies

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 cell 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

Cooperative ordering of treadmilling filaments in cytoskeletal networks of FtsZ and its crosslinker ZapA

During bacterial cell division, the tubulin-homolog FtsZ forms a ring-like structure at the center of the cell. This Z-ring not only organizes the division machinery, but treadmilling of FtsZ filaments was also found to play a key role in distributing proteins at the division site. What regulates the architecture, dynamics and stability of the Z-ring is currently unknown, but FtsZ-associated proteins are known to play an important role. Here, using an in vitro reconstitution approach, we studied how the well-conserved protein ZapA affects FtsZ treadmilling and filament organization into large-scale patterns. Using high-resolution fluorescence microscopy and quantitative image analysis, we found that ZapA cooperatively increases the spatial order of the filament network, but binds only transiently to FtsZ filaments and has no effect on filament length and treadmilling velocity. Together, our data provides a model for how FtsZ-associated proteins can increase the precision and stability of the bacterial cell division machinery in a switch-like manner

Project MOSI: rationale and pilot-study results of an initiative to help protect zoo animals from mosquito-transmitted pathogens and contribute data on mosquito spatio–temporal distribution change

Mosquito-borne pathogens pose major threats to both wildlife and human health and, largely as a result of unintentional human-aided dispersal of their vector species, their cumulative threat is on the rise. Anthropogenic climate change is expected to be an increasingly significant driver of mosquito dispersal and associated disease spread. The potential health implications of changes in the spatio-temporal distribution of mosquitoes highlight the importance of ongoing surveillance and, where necessary, vector control and other health-management measures. The World Association of Zoos and Aquariums initiative, Project MOSI, was established to help protect vulnerable wildlife species in zoological facilities from mosquito-transmitted pathogens by establishing a zoo-based network of fixed mosquito monitoring sites to assist wildlife health management and contribute data on mosquito spatio-temporal distribution changes. A pilot study for Project MOSI is described here, including project rationale and results that confirm the feasibility of conducting basic standardized year-round mosquito trapping and monitoring in a zoo environment