Mauro Mobilia

Mobility promotes and jeopardizes biodiversity in rock–paper–scissors games

Biodiversity is essential to the viability of ecological systems. Species diversity in ecosystems is promoted by cyclic, non-hierarchical interactions among competing populations. Central features of such non-transitive relations are represented by the ‘rock–paper–scissors’ game, in which rock crushes scissors, scissors cut paper, and paper wraps rock. In combination with spatial dispersal of static populations, this type of competition results in the stable coexistence of all species and the long-term maintenance of biodiversity. However, population mobility is a central feature of real ecosystems: animals migrate, bacteria run and tumble. Here, we observe a critical influence of mobility on species diversity. When mobility exceeds a certain value, biodiversity is jeopardized and lost. In contrast, below this critical threshold all subpopulations coexist and an entanglement of travelling spiral waves forms in the course of time. We establish that this phenomenon is robust; it does not depend on the details of cyclic competition or spatial environment. These findings have important implications for maintenance and temporal development of ecological systems and are relevant for the formation and propagation of patterns in microbial populations or excitable media.

Cyclic dominance in evolutionary games: A review

Rock is wrapped by paper, paper is cut by scissors and scissors are crushed by rock. This simple game is popular among children and adults to decide on trivial disputes that have no obvious winner, but cyclic dominance is also at the heart of predator–prey interactions, the mating strategy of side-blotched lizards, the overgrowth of marine sessile organisms and competition in microbial populations. Cyclical interactions also emerge spontaneously in evolutionary games entailing volunteering, reward, punishment, and in fact are common when the competing strategies are three or more, regardless of the particularities of the game. Here, we review recent advances on the rock–paper–scissors (RPS) and related evolutionary games, focusing, in particular, on pattern formation, the impact of mobility and the spontaneous emergence of cyclic dominance. We also review mean-field and zero-dimensional RPS models and the application of the complex Ginzburg–Landau equation, and we highlight the importance and usefulness of statistical physics for the successful study of large-scale ecological systems. Directions for future research, related, for example, to dynamical effects of coevolutionary rules and invasion reversals owing to multi-point interactions, are also outlined.

On the role of zealotry in the voter model

We study the voter model with a finite density of zealots—voters that never change opinion. For equal numbers of zealots of each species, the distribution of magnetization (opinions) is Gaussian in the mean-field limit, as well as in one and two dimensions, with a width that is proportional to , where Z is the number of zealots, independent of the total number of voters. Thus just a few zealots can prevent consensus or even the formation of a robust majority.

Noise and correlations in a spatial population model with cyclic competition.

Noise and spatial degrees of freedom characterize most ecosystems. Some aspects of their influence on the coevolution of populations with cyclic interspecies competition have been demonstrated in recent experiments [e.g., B. Kerr, Nature (London) 418, 171 (2002)10.1038/nature00823]. To reach a better theoretical understanding of these phenomena, we consider a paradigmatic spatial model where three species exhibit cyclic dominance. Using an individual-based description, as well as stochastic partial differential and deterministic reaction-diffusion equations, we account for stochastic fluctuations and spatial diffusion at different levels and show how fascinating patterns of entangled spirals emerge. We rationalize our analysis by computing the spatiotemporal correlation functions and provide analytical expressions for the front velocity and the wavelength of the propagating spiral waves.

Majority versus minority dynamics: Phase transition in an interacting two-state spin system

We introduce a simple model of opinion dynamics in which binary-state agents evolve due to the influence of agents in a local neighborhood. In a single update step, a fixed-size group is defined and all agents in the group adopt the state of the local majority with probability p or that of the local minority with probability 1-p. For group size G=3, there is a phase transition at p(c)=2/3 in all spatial dimensions. For p>p(c), the global majority quickly predominates, while for p<p(c), the system is driven to a mixed state in which the densities of agents in each state are equal. For p=p(c), the average magnetization (the difference in the density of agents in the two states) is conserved and the system obeys classical voter model dynamics. In one dimension and within a Kirkwood decoupling scheme, the final magnetization in a finite-length system has a nontrivial dependence on the initial magnetization for all p not equal p(c), in agreement with numerical results. At p(c), the exact two-spin correlation functions decay algebraically toward the value 1 and the system coarsens as in the classical voter model.

Phase Transitions and Spatio-Temporal Fluctuations in Stochastic Lattice Lotka–Volterra Models

We study the general properties of stochastic two-species models for predator-prey competition and coexistence with Lotka–Volterra type interactions defined on a d-dimensional lattice. Introducing spatial degrees of freedom and allowing for stochastic fluctuations generically invalidates the classical, deterministic mean-field picture. Already within mean-field theory, however, spatial constraints, modeling locally limited resources, lead to the emergence of a continuous active-to-absorbing state phase transition. Field-theoretic arguments, supported by Monte Carlo simulation results, indicate that this transition, which represents an extinction threshold for the predator population, is governed by the directed percolation universality class. In the active state, where predators and prey coexist, the classical center singularities with associated population cycles are replaced by either nodes or foci. In the vicinity of the stable nodes, the system is characterized by essentially stationary localized clusters of predators in a sea of prey. Near the stable foci, however, the stochastic lattice Lotka–Volterra system displays complex, correlated spatio-temporal patterns of competing activity fronts. Correspondingly, the population densities in our numerical simulations turn out to oscillate irregularly in time, with amplitudes that tend to zero in the thermodynamic limit. Yet in finite systems these oscillatory fluctuations are quite persistent, and their features are determined by the intrinsic interaction rates rather than the initial conditions. We emphasize the robustness of this scenario with respect to various model perturbations.

Oscillatory dynamics in rock–paper–scissors games with mutations

We study the oscillatory dynamics in the generic three-species rock-paper-scissors games with mutations. In the mean-field limit, different behaviors are found: (a) for high mutation rate, there is a stable interior fixed point with coexistence of all species; (b) for low mutation rates, there is a region of the parameter space characterized by a limit cycle resulting from a Hopf bifurcation; (c) in the absence of mutations, there is a region where heteroclinic cycles yield oscillations of large amplitude (not robust against noise). After a discussion on the main properties of the mean-field dynamics, we investigate the stochastic version of the model within an individual-based formulation. Demographic fluctuations are therefore naturally accounted and their effects are studied using a diffusion theory complemented by numerical simulations. It is thus shown that persistent erratic oscillations (quasi-cycles) of large amplitude emerge from a noise-induced resonance phenomenon. We also analytically and numerically compute the average escape time necessary to reach a (quasi-)cycle on which the system oscillates at a given amplitude.

Bottleneck-induced transitions in a minimal model for intracellular transport

We consider the influence of disorder on the nonequilibrium steady state of a minimal model for intracellular transport. In this model particles move unidirectionally according to the totally asymmetric exclusion process (TASEP) and are coupled to a bulk reservoir by Langmuir kinetics. Our discussion focuses on localized point defects acting as a bottleneck for the particle transport. Combining analytic methods and numerical simulations, we identify a rich phase behavior as a function of the defect strength. Our analytical approach relies on an effective mean-field theory obtained by splitting the lattice into two subsystems, which are effectively connected exploiting the local current conservation. Introducing the key concept of a carrying capacity, the maximal current that can flow through the bulk of the system (including the defect), we discriminate between the cases where the defect is irrelevant and those where it acts as a bottleneck and induces various novel phases (called bottleneck phases). Contrary to the simple TASEP in the presence of inhomogeneities, many scenarios emerge and translate into rich underlying phase diagrams, the topological properties of which are discussed.

Fluctuations and correlations in lattice models for predator-prey interaction.

Including spatial structure and stochastic noise invalidates the classical Lotka-Volterra picture of stable regular population cycles emerging in models for predator-prey interactions. Growth-limiting terms for the prey induce a continuous extinction threshold for the predator population whose critical properties are in the directed percolation universality class. We discuss the robustness of this scenario by considering an ecologically inspired stochastic lattice predator-prey model variant where the predation process includes next-nearest-neighbor interactions. We find that the corresponding stochastic model reproduces the above scenario in dimensions 1< d < or =4, in contrast with the mean-field theory, which predicts a first-order phase transition. However, the mean-field features are recovered upon allowing for nearest-neighbor particle exchange processes, provided these are sufficiently fast.

Spatial rock-paper-scissors models with inhomogeneous reaction rates

We study several variants of the stochastic four-state rock-paper-scissors game or, equivalently, cyclic three-species predator-prey models with conserved total particle density, by means of Monte Carlo simulations on one- and two-dimensional lattices. Specifically, we investigate the influence of spatial variability of the reaction rates and site occupancy restrictions on the transient oscillations of the species densities and on spatial correlation functions in the quasistationary coexistence state. For small systems, we also numerically determine the dependence of typical extinction times on the number of lattice sites. In stark contrast with two-species stochastic Lotka-Volterra systems, we find that for our three-species models with cyclic competition quenched disorder in the reaction rates has very little effect on the dynamics and the long-time properties of the coexistence state. Similarly, we observe that site restriction only has a minor influence on the systems dynamical properties. Our results therefore demonstrate that the features of the spatial rock-paper-scissors system are remarkably robust with respect to model variations, and stochastic fluctuations as well as spatial correlations play a comparatively minor role.