This paper presents a review and critical analysis of the mathematical literature concerning the modeling of vehicular traffic and crowd phenomena. The survey of models deals with the representation scales and the mathematical frameworks that are used for the modeling approach. The paper also considers the challenging objective of modeling complex systems consisting of large systems of individuals interacting in a nonlinear manner, where one of the modeling difficulties is the fact that these systems are difficult to model at a global level when based only on the description of the dynamics of individual elements. The review is concluded with a critical analysis focused on research perspectives that consider the development of a unified modeling strategy.

On the Modeling of Traffic and Crowds: A Survey of Models, Speculations, and Perspectives

Self-organization in developing living organisms relies on the capability of cells to duplicate and perform a collective motion inside the surrounding environment. Chemical and mechanical interactions coordinate such a cooperative behaviour, driving the dynamical evolution of the macroscopic system. In this work, we perform an analytical and computational analysis to study pattern formation during the spreading of an initially circular bacterial colony on a Petri dish. The continuous mathematical model addresses the growth and the chemotactic migration of the living monolayer, together with the diffusion and consumption of nutrients in the agar. The governing equations contain four dimensionless parameters, accounting for the interplay among the chemotactic response, the bacteria–substrate interaction and the experimental geometry. The spreading colony is found to be always linearly unstable to perturbations of the interface, whereas branching instability arises in finite-element numerical simulations. The typical length scales of such fingers, which align in the radial direction and later undergo further branching, are controlled by the size parameters of the problem, whereas the emergence of branching is favoured if the diffusion is dominant on the chemotaxis. The model is able to predict the experimental morphologies, confirming that compact (resp. branched) patterns arise for fast (resp. slow) expanding colonies. Such results, while providing new insights into pattern selection in bacterial colonies, may finally have important applications for designing controlled patterns.

https://royalsocietypublishing.org/doi/pdf/10.1098/rsif.2014.1290

Branching instability in expanding bacterial colonies

How can repulsive and attractive forces, acting on a conservative system, create stable traveling patterns or branching instabilities? We have proposed to study this question in the framework of the hyperbolic Keller-Segel system with logistic sensitivity. This is a model system motivated by experiments on cell communities auto-organization, a field which is also called socio-biology. We continue earlier modeling work, where we have shown numerically that branching patterns arise for this system and we have analyzed this instability by formal asymptotics for small diffusivity of the chemo-repellent. Here we are interested in the more general situation, where the diffusivities of both the chemo-attractant and the chemo-repellent are positive. To do so, we develop an appropriate functional analysis framework. We apply our method to two cases. Firstly we analyze steady states. Secondly we analyze traveling waves when neglecting the degradation coefficient of the chemo-repellent; the unique wave speed appears through a singularity cancelation which is the main theoretical difficulty. This shows that in different situations the cell density takes the shape of a plateau. The existence of steady states and traveling plateaus are a symptom of how rich the system is and why branching instabilities can occur. Numerical tests show that large plateaus may split into smaller ones, which remain stable.

Traveling plateaus for a hyperbolic Keller-Segel system with attraction and repulsion: existence and branching instabilities

In order to observe growth phenomena in biology where dendritic shapes appear, we propose a simple model where a given population evolves feeded by a diffusing nutriment, but is subject to a density constraint. The particles (e.g., cells) of the population spontaneously stay passive at rest, and only move in order to satisfy the constraint $\rho\leq 1$, by choosing the minimal correction velocity so as to prevent overcongestion. We treat this constraint by means of projections in the space of densities endowed with the Wasserstein distance $W_2$, defined through optimal transport. This allows to provide an existence result and suggests some numerical computations, in the same spirit of what the authors did for crowd motion (but with extra difficulties, essentially due to the fact that the total mass may increase). The numerical simulations show, according to the values of the parameter and in particular of the diffusion coefficient of the nutriment, the formation of dendritic patterns in the space occupied by cells.

Congestion-driven dendritic growth

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 instability of the circular colony shape and the subsequent formation of fingers. In this work, we study the effect of bacterial surfactant production and substrate wettability on colony growth and shape within the framework of a hydrodynamic thin film model. We show that variations in the wettability and surfactant production are sufficient to reproduce four different types of colony growth, which have been described in the literature, namely, arrested and continuous spreading of circular colonies, slightly modulated front lines and the formation of pronounced fingers.

Modelling of surfactant-driven front instabilities in spreading bacterial colonies.

Bacillus subtilis swarms rapidly over the surface of a synthetic medium creating remarkable hyperbranched dendritic communities. Models to reproduce such effects have been proposed under the form of parabolic Partial Differential Equations representing the dynamics of the active cells (both motile and multiplying), the passive cells (non-motile and non-growing) and nutrient concentration. We test the numerical behavior of such models and compare them to relevant experimental data together with a critical analysis of the validity of the models based on recent observations of the swarming bacteria which show that nutrients are not limitating but distinct subpopulations growing at different rates are likely present.

/pdf/models-of-self-organizing-bacterial-communities-and-4byk95k5v3.pdf

I. B. Holland

Papers

Models of Self-Organizing Bacterial Communities and Comparisons with Experimental Observations