Many bacterial species exchange signaling molecules to coordinate population-wide responses. For this process known as quorum sensing the concentration of the respective molecules is crucial. Here we consider the interaction between spatially distributed bacterial colonies so that the spreading of the signaling molecules in space becomes important. The exponential growth of the signal-producing populations and the corresponding increase in signaling molecule production result in an exponential concentration profile that spreads with uniform speed. The theoretical predictions are supported by experiments with different strains of the soil bacterium Sinorhizobium meliloti that display fluorescence when either producing or responding to the signaling molecules.
Colonies of bacteria grown on thin agar plate exhibit fractal patterns as a result of adaptation to their environments. The bacterial colony pattern formation is regulated crucially by chemotaxis, the movement of cells along a chemical concentration gradient. Here, the dynamics of pattern formation in bacterial colony is investigated theoretically through a continuum model that considers chemotaxis. In the case of the gradient sensed by the bacterium is nearly uniform, the bacterial colony patterns are self-similar, which they look the same at every scale. The scaling law of the bacterial colo…
The spreading of bacterial colonies at solid-air interfaces is determined by the physico-chemical properties of the involved interfaces. The production of surfactant molecules by bacteria is a widespread strategy that allows the colony to efficiently expand over the substrate. On the one hand, surfactant molecules lower the surface tension of the colony, effectively increasing the wettability of the substrate, which facilitates spreading. On the other hand, gradients in the surface concentration of surfactant molecules result in Marangoni flows that drive spreading. These flows may cause an in…
Bacterial conglomerates such as biofilms and microcolonies are ubiquitous in nature and play an important role in industry and medicine. In contrast to well-mixed, diluted cultures routinely used in microbial research, bacteria in a microcolony interact mechanically with one another and with the substrate to which they are attached. Despite their ubiquity, little is known about the role of such mechanical interactions on growth and biological evolution of microbial populations. Here we use a computer model of a microbial colony of rod-shaped cells to investigate how physical interactions betwe…
Colonies of rod-shaped bacteria constitute a system of colloidal active matter with nematic properties. As a single initial bacterium multiplies through repeated divisions, the resulting colony quickly loses long-range orientational order, but retains locally ordered domains. At the boundaries of these domains, topological defects emerge, which move around randomly as the colony grows. In both experiments and simulations, we find that these defects are created at a rate that corresponds to the exponential growth of the colony, resulting in a stable defect density. Both this defect density and …
This paper introduces a reaction-diffusion-chemotaxis model for bacterial aggregation patterns on the surface of thin agar plates. It is based on the non-linear degenerate cross diffusion model proposed by Kawasaki et al. (J. of Theor. Biol. 188(2) 1997) and it includes a suitable nutrient chemotactic term compatible with such type of diffusion. High resolution numerical simulations using Graphic Processing Units (GPUs) of the new model are presented, showing that the chemotactic term enhances the velocity of propagation of the colony envelope for dense-branching morphologies. In addition, the…
Bacteria commonly live in structured communities that affect human health and influence ecological systems. Heterogeneous populations, such as motile and non-motile populations, often coexist in bacteria communities. Motile subpopulations in microbial communities are believed to be important to dispersal, quest for food, and material transport. However, except in circumstances where motile cells drive colony expansion (e.g. bacterial swarming), the physiological functions of motile subpopulations in bacterial communities are largely unclear. Here we discovered that motile cells in routinely cu…
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 mic…
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 …
It is widely accepted that population genetics theory is the cornerstone of evolutionary analyses. Empirical tests of the theory, however, are challenging because of the complex relationships between space, dispersal, and evolution. Critically, we lack quantitative validation of the spatial models of population genetics. Here we combine analytics, on and off-lattice simulations, and experiments with bacteria to perform quantitative tests of the theory. We study two bacterial species, the gut microbe Escherichia coli and the opportunistic pathogen Pseudomonas aeruginosa, and show that spatio-ge…
Rod-shaped bacteria, such as Escherichia coli, commonly live forming mounded colonies. They initially grow two-dimensionally on a surface and finally achieve three-dimensional growth. While it was recently reported that three-dimensional growth is promoted by topological defects of winding number $+1/2$ in populations of motile bacteria, how cellular alignment plays a role in non-motile cases is largely unknown. Here, we investigate the relevance of topological defects in colony formation processes of non-motile E. coli populations, and found that both $\pm 1/2$ topological defects contribute …
It has been argued that there is biological and modeling evidence that a non-linear diffusion coefficient of the type D(b) = D_0 b^{k} underlies the formation of a number of growth patterns of bacterial colonies. We study a reaction-diffusion system with a non-linear diffusion coefficient introduced by Ben-Jacob et al. Due to the fact that the bacterial diffusion coefficient vanishes when the bacterial density b -> 0, the standard linear stability analysis for fronts cannot be used. We introduce an extension of the stability analysis which can be applied to such singular fronts, map out the re…
The effect of mechanical interactions between cells in the spreading of bacterial populations was investigated in one-dimensional space. A continuum-mechanics approach, comprising cell migration, proliferation, and exclusion processes, was employed to elucidate the dynamics. The consequent nonlinear reaction-diffusion-like equation describes the constitution dynamics of a bacterial population. In this model, bacterial cells were treated as rod-like particles that interact with each other through hard-core repulsion, which introduces the exclusion effect that causes bacterial populations to mig…
CRISPR-Cas mediated immunity in bacteria allows bacterial populations to protect themselves against pathogens. However, it also exposes them to the dangers of auto-immunity by developing protection that targets its own genome. Using a simple model of the coupled dynamics of phage and bacterial populations, we explore how acquisition rates affect the survival rate of the bacterial colony. We find that the optimal strategy depends on the initial population sizes of both viruses and bacteria. Additionally, certain combinations of acquisition and dynamical rates and initial population sizes guaran…
Directed Percolation (DP) is a classic model for nonequilibrium phase transitions into a single absorbing state (fixation). It has been extensively studied by analytical and numerical techniques in diverse contexts. Recently, DP has appeared as a generic model for the evolutionary/ecological dynamics of competing bacterial populations. Range expansion -- the stochastic reproduction of bacteria competing for space to be occupied by their progeny -- leads to a fluctuating and rough growth front, which is known from experiment and simulation to affect the underlying critical behavior of the DP tr…
We study the spreading of a bacterial colony undergoing turbulent like collective motion. We present two minimalistic models to investigate the interplay between population growth and coherent structures arising from turbulence. Using Direct Numerical Simulation (DNS) of the proposed models we find that turbulence has two prominent effects on the spatial growth of the colony: (a) the front speed is enhanced, and (b) the front gets crumpled. Both these effects, which we highlight by using statistical tools, are markedly different in our two models. We also show that the crumpled front structure…
In nature, bacterial collectives typically consist of multiple species, which are interacting both biochemically and physically. Nonetheless, past studies on the physical properties of swarming bacteria were focused on axenic (single species) populations. In bacterial swarming, intricate interactions between the individuals lead to clusters, rapid jets and vortices that depend on cell characteristics such as speed and length. In this work, we show the first results of rapidly swarming mixed-species populations of {\sl Bacillus subtilis} and {\sl Serratia marcescens}, two model swarm species th…
The expansion of biological species in natural environments is usually described as the combined effect of individual spatial dispersal and growth. In the case of aquatic ecosystems flow transport can also be extremely relevant as an extra, advection induced, dispersal factor. There is a lack of reproducible experimental studies on biological fronts of living organisms in controlled streaming habitats. It is thus not clear if, and to which extent, the current theoretical and experimental knowledge on advective-reactive-diffusive fronts for chemical reactions can also apply to the expansion of …
We propose a model for bacterial Quorum Sensing based on an auxiliary electrostatic-like interac-tion originating from a fictitious electrical charge that represents bacteria activity. A cooperative mechanism for charge/activity exchange is introduced to implement chemotaxis and replication. The bacteria system is thus represented by means of a complex resistor network where link re-sistances take into account the allowed activity-flow among individuals. By explicit spatial sto-chastic simulations, we show that the model exhibits different quasi-realistic behaviors from colo-ny formation to bi…
We observe that small drops of a Bacillus subtilis suspension deposited on agar strongly increase in volume while similar bacteria-void drops do not. By measuring the bacterial concentration within the drop at different heights, we show that the biomass increase due to the constant bacterial cell-division is too small to explain the drop bloating. Rather, the increased volume is caused by the presence of surfactin - a surfactant produced by the bacteria - which induces a water flow out of the environment by an osmotic capillary effect. The required concentration is very low (< 1 mM), four orde…
The spreading of bacterial populations is central to processes in agriculture, the environment, and medicine. However, existing models of spreading typically focus on cells in unconfined settings--despite the fact that many bacteria inhabit complex and crowded environments, such as soils, sediments, and biological tissues/gels, in which solid obstacles confine the cells and thereby strongly regulate population spreading. Here, we develop an extended version of the classic Keller-Segel model of bacterial spreading that incorporates the influence of confinement in promoting both cell-solid and c…
Sedimentation in active fluids has come into focus due to the ubiquity of swimming micro-organisms in natural and artificial environments. Here, we experimentally investigate sedimentation of passive particles in water containing various concentrations of the bacterium E. coli. Results show that the presence of living bacteria reduces the velocity of the sedimentation front even in the dilute regime, where the sedimentation velocity is expected to be independent of particle concentration. Bacteria increase the effective diffusion coefficient of the passive particles, which determines the width…
We study the migration of chemotactic wild-type Escherichia coli populations in semisolid (soft) agar in the concentration range C = 0.15-0.5% (w/v). For C < 0.35%, expanding bacterial colonies display characteristic chemotactic rings. At C = 0.35%, however, bacteria migrate as broad circular bands rather than sharp rings. These are growth/diffusion waves arising because of suppression of chemotaxis by the agar and have not been previously reported experimentally to our knowledge. For C = 0.4-0.5%, expanding colonies do not span the depth of the agar and develop pronounced front instabilities.…
