Bacterial swarming is a rapid mass-migration, in which thousands of cells spread collectively to colonize a surface. Physically, swarming is a natural example of active particles that use energy to generate motion. Accordingly, understanding the constraints physics imposes on the dynamics is essential to understand the mechanisms underlying the swarming phenomenon. We present new experiments of swarming Bacillus subtilis mutants with different aspect ratios and densities. Analyzing the dynamics reveals a rich phase diagram of qualitatively distinct swarming regimes, describing how the shape and density of cells govern the global dynamical characteristics of the entire swarm. Moreover, 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. This contrasts characteristic clustering behavior of selfpropelled rods that is recovered only for very elongated mutant species. Thus, bacteria have adapted their physics to optimize the principle functions assumed for swarming. 2 Micro-organisms such as bacteria, sperm-, epithelial-and cancer-cells, as well as immune T-cells and even inanimate active particles, generate collective flows and demonstrate a wealth of newly discovered emergent dynamical patterns [Szabó Be'er 2019]. This report addresses the dynamics of swarming bacteria a biological state to which some bacterial species transition, in which rod-shaped cells, powered by flagellar rotation, migrate rapidly on surfaces en-mass [Harshey 2003; Darnton 2010; Kearns 2010; Tuson 2013]. Swarming allows efficient expansion and colonization of new territories, even under harsh and adverse conditions such as starvation or antibiotic stress [Lai 2009; Benisty 2015]. Revealing the biological and physical mechanisms underlying bacterial swarming is therefore a key to our understanding of how bacteria spread and invade new niches. The transition to swarming involves several critical intra-cellular processes such as an increase in flagellar number and changes in cell shape, suggesting these changes promote favorable swarming conditions [Harshey 2003; Kearns 2010; Tuson 2013; Mukherjee 2015; Be'er 2019]. Quantifying the "quality" of swarming can be done using the tools of statistical physics by analyzing the dynamical properties of large, out-ofequilibrium, self-propelled collectives [Ramaswamy 2010; Vicsek 2012; Bär 2019].Accordingly, one of the primary goals of such quantification is to obtain a phase diagram that would describe the possible dynamical states of swarms as a function of controlled parameters such as density and cell aspect ratio. As bacteria move and grow, they trace an effective path through the phase diagram, transitioning between different swarming regimes. Thus, a phase diagram provides a map that explains how the microscopic mechanical properties of cells, which are regulated by complex bio-
Swarming bacteria collectively migrate on surfaces using flagella, forming dynamic whirls and jets that consist of millions of individuals. Because some swarming bacteria elongate prior to actual motion, cell aspect ratio may play a significant role in the collective dynamics. Extensive research on self-propelled rod-like particles confirms that elongation promotes alignment, strongly affecting the dynamics. Here, we study experimentally the collective dynamics of variants of swarming Bacillus subtilis, that differ in length. We show that the swarming statistics depends on aspect ratio in a critical, fundamental fashion not predicted by theory. The fastest motion was obtained for the wildtype and variants that are similar in length. However, shorter and longer cells exhibit anomalous, non-Gaussian statistics and non-exponential decay of the auto-correlation function, indicating lower collective motility. These results suggest that the robust mechanisms to maintain aspect ratios may be important for efficient swarming motility. Wild-type cells are optimal in this sense.
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