Individual swimming bacteria are known to bias their random trajectories in search of food and to optimize survival. The motion of bacteria within a swarm, wherein they migrate as a collective group over a solid surface, is fundamentally different as typical bacterial swarms show large-scale swirling and streaming motions involving millions to billions of cells. Here by tracking trajectories of fluorescently labelled individuals within such dense swarms, we find that the bacteria are performing super-diffusion, consistent with Lévy walks. Lévy walks are characterized by trajectories that have straight stretches for extended lengths whose variance is infinite. The evidence of super-diffusion consistent with Lévy walks in bacteria suggests that this strategy may have evolved considerably earlier than previously thought.

Over the past decade, technological advances in experimental and animal tracking techniques have motivated a renewed theoretical interest in animal collective motion and, in particular, locust swarming. This review offers a comprehensive biological background followed by comparative analysis of recent models of locust collective motion, in particular locust marching, their settings, and underlying assumptions. We describe a wide range of recent modeling and simulation approaches, from discrete agent-based models of self-propelled particles to continuous models of integro-differential equations, aimed at describing and analyzing the fascinating phenomenon of locust collective motion. These modeling efforts have a dual role: The first views locusts as a quintessential example of animal collective motion. As such, they aim at abstraction and coarse-graining, often utilizing the tools of statistical physics. The second, which originates from a more biological perspective, views locust swarming as a scientific problem of its own exceptional merit. The main goal should, thus, be the analysis and prediction of natural swarm dynamics. We discuss the properties of swarm dynamics using the tools of statistical physics, as well as the implications for laboratory experiments and natural swarms. Finally, we stress the importance of a combined-interdisciplinary, biological-theoretical effort in successfully confronting the challenges that locusts pose at both the theoretical and practical levels.

Bacterial swarming is a collective mode of motion in which cells migrate rapidly over surfaces, forming dynamic patterns of whirls and jets. This review presents a physical point of view of swarming bacteria, with an emphasis on the statistical properties of the swarm dynamics as observed in experiments. The basic physical principles underlying the swarm and their relation to contemporary theories of collective motion and active matter are reviewed and discussed in the context of the biological properties of swarming cells. We suggest a paradigm according to which bacteria have optimized some of their physical properties as a strategy for rapid surface translocation. In other words, cells take advantage of favorable physics, enabling efficient expansion that enhances survival under harsh conditions.

Bacterial swarming is a rapid mass-migration, in which thousands of cells spread collectively to colonize surfaces. Physically, swarming is a natural example for active particles that use energy to generate motion. Accordingly, understanding the constraints physics imposes on these dynamics is essential for understanding the mechanisms underlying swarming. We present new experiments of swarming Bacillus subtilis mutants with different aspect ratios and at different densities; two physical quantities known to be associated with collective behavior. Analyzing the dynamics reveals a rich phase diagram of qualitatively distinct swarming regimes, describing how cell shape and population density govern the dynamical characteristics of the swarm. In particular, we show that under standard conditions, bacteria inhabit a region of phase space that is associated with rapid mixing and robust dynamics, with homogeneous density and no preferred direction of motion. The results suggest that bacteria have adapted their physical properties to optimize the principle functions assumed for swarming.

The principal interactions leading to the emergence of order in swarms of marching locust nymphs was studied both experimentally, using small groups of marching locusts in the lab, and using computer simulations. We utilized a custom tracking algorithm to reveal fundamental animal-animal interactions leading to collective motion. Uncovering this behavior introduced a new agent-based modeling approach in which pause-and-go motion is pivotal. The behavioral and modeling findings are largely based on motion-related visual sensory inputs obtained by the individual locust. Results suggest a generic principle, in which intermittent animal motion can be considered as a sequence of individual decisions as animals repeatedly reassess their situation and decide whether or not to swarm. This interpretation implies, among other things, some generic characteristics regarding the build-up and emergence of collective order in swarms: in particular, that order and disorder are generic meta-stable states of the system, suggesting that the emergence of order is kinetic and does not necessarily require external environmental changes. This work calls for further experimental as well as theoretical investigation of the neural mechanisms underlying locust coordinative behavior.

Bacteria organize in a variety of collective states, from swarming—rapid surface exploration, to biofilms—highly dense immobile communities attributed to stress resistance. It has been suggested that biofilm and swarming are oppositely controlled, making this transition particularly interesting for understanding the ability of bacterial colonies to adapt to challenging environments. Here, the swarm to biofilm transition is studied in Bacillus subtilis by analyzing the bacterial dynamics both on the individual and collective scales. We show that both biological and physical processes facilitate the transition. A few individual cells that initiate the biofilm program cause nucleation of large, approximately scale-free, stationary aggregates of trapped swarm cells. Around aggregates, cells continue swarming almost unobstructed, while inside, trapped cells are added to the biofilm. While our experimental findings rule out previously suggested purely physical effects as a trigger for biofilm formation, they show how physical processes, such as clustering and jamming, accelerate biofilm formation.