Bacteria form human and animal microbiota. They are the leading causes of many infections and constitute an important class of active matter. Concentrated bacterial suspensions exhibit large-scale turbulent-like locomotion and swarming. While the collective behavior of bacteria in Newtonian fluids is relatively well understood, many fundamental questions remain open for complex fluids. Here, we report on the collective bacterial motion in a representative biological non-Newtonian viscoelastic environment exemplified by mucus. Experiments are performed with synthetic porcine gastric mucus, natural cow cervical mucus, and a Newtonian-like polymer solution. We have found that an increase in mucin concentration and, correspondingly, an increase in the suspension’s elasticity monotonously increases the length scale of collective bacterial locomotion. On the contrary, this length remains practically unchanged in Newtonian polymer solution in a wide range of concentrations. The experimental observations are supported by computational modeling. Our results provide insight into how viscoelasticity affects the spatiotemporal organization of bacterial active matter. They also expand our understanding of bacterial colonization of mucosal surfaces and the onset of antibiotic resistance due to swarming.
Bacterial active matter
Abstract Bacteria are among the oldest and most abundant species on Earth. Bacteria successfully colonize diverse habitats and play a significant role in the oxygen, carbon, and nitrogen cycles. They also form human and animal microbiota and may become sources of pathogens and a cause of many infectious diseases. Suspensions of motile bacteria constitute one of the most studied examples of active matter: a broad class of non-equilibrium systems converting energy from the environment (e.g., chemical energy of the nutrient) into mechanical motion. Concentrated bacterial suspensions, often termed active fluids, exhibit complex collective behavior, such as large-scale turbulent-like motion (so-called bacterial turbulence) and swarming. The activity of bacteria also affects the effective viscosity and diffusivity of the suspension. This work reports on the progress in bacterial active matter from the physics viewpoint. It covers the key experimental results, provides a critical assessment of major theoretical approaches, and addresses the effects of visco-elasticity, liquid crystallinity, and external confinement on collective behavior in bacterial suspensions.
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- Award ID(s):
- 2140010
- PAR ID:
- 10427974
- Date Published:
- Journal Name:
- Reports on Progress in Physics
- Volume:
- 85
- Issue:
- 7
- ISSN:
- 0034-4885
- Page Range / eLocation ID:
- 076601
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract -
As a paradigmatic model of active fluids, bacterial suspensions show intriguing rheological responses drastically different from their counterpart colloidal suspensions. Although the flow of bulk bacterial suspensions has been extensively studied, the rheology of bacterial suspensions under confinement has not been experimentally explored. Here, using a microfluidic viscometer, we systematically measure the rheology of dilute Escherichia coli suspensions under different degrees of confinement. Our study reveals a strong confinement effect: the viscosity of bacterial suspensions decreases substantially when the confinement scale is comparable or smaller than the run length of bacteria. Moreover, we also investigate the microscopic dynamics of bacterial suspensions including velocity profiles, bacterial density distributions, and single bacterial dynamics in shear flows. These measurements allow us to construct a simple heuristic model based on the boundary layer of upstream swimming bacteria near confining walls, which qualitatively explains our experimental observations. Our study sheds light on the influence of the boundary layer of collective bacterial motions on the flow of confined bacterial suspensions. Our results provide a benchmark for testing different rheological models of active fluids and are useful for understanding the transport of microorganisms in confined geometries.more » « less
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Drying of bacterial suspensions is frequently encountered in a plethora of natural and engineering processes. However, the evaporation-driven mechanical instabilities of dense consolidating bacterial suspensions have not been explored heretofore. Here, we report the formation of two different crack patterns of drying suspensions of Escherichia coli ( E. coli ) with distinct motile behaviors. Circular cracks are observed for wild-type E. coli with active swimming, whereas spiral-like cracks form for immotile bacteria. Using the elastic fracture mechanics and the poroelastic theory, we show that the formation of the circular cracks is determined by the tensile nature of the radial drying stress once the cracks are initiated by the local order structure of bacteria due to their collective swimming. Our study demonstrates the link between the microscopic swimming behaviors of individual bacteria and the mechanical instabilities and macroscopic pattern formation of drying bacterial films. The results shed light on the dynamics of active matter in a drying process and provide useful information for understanding various biological processes associated with drying bacterial suspensions.more » « less
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Bacterial suspensions—a premier example of active fluids—show an unusual response to shear stresses. Instead of increasing the viscosity of the suspending fluid, the emergent collective motions of swimming bacteria can turn a suspension into a superfluid with zero apparent viscosity. Although the existence of active superfluids has been demonstrated in bulk rheological measurements, the microscopic origin and dynamics of such an exotic phase have not been experimentally probed. Here, using high-speed confocal rheometry, we study the dynamics of concentrated bacterial suspensions under simple planar shear. We find that bacterial superfluids under shear exhibit unusual symmetric shear bands, defying the conventional wisdom on shear banding of complex fluids, where the formation of steady shear bands necessarily breaks the symmetry of unsheared samples. We propose a simple hydrodynamic model based on the local stress balance and the ergodic sampling of nonequilibrium shear configurations, which quantitatively describes the observed symmetric shear-banding structure. The model also successfully predicts various interesting features of swarming vortices in stationary bacterial suspensions. Our study provides insights into the physical properties of collective swarming in active fluids and illustrates their profound influences on transport processes.more » « less
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null (Ed.)We experimentally study the emergence of collective bacterial swimming, a phenomenon often referred to as bacterial turbulence. A phase diagram of the flow of 3D Escherichia coli suspensions spanned by bacterial concentration, the swimming speed of bacteria, and the number fraction of active swimmers is systematically mapped, which shows quantitative agreement with kinetic theories and demonstrates the dominant role of hydrodynamic interactions in bacterial collective swimming. We trigger bacterial turbulence by suddenly increasing the swimming speed of light-powered bacteria and image the transition to the turbulence in real time. Our experiments identify two unusual kinetic pathways, i.e., the one-step transition with long incubation periods near the phase boundary and the two-step transition driven by long-wavelength instabilities deep inside the turbulent phase. Our study provides not only a quantitative verification of existing theories but also insights into interparticle interactions and transition kinetics of bacterial turbulence.more » « less
- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1088/1361-6633/ac723d
