Ordering dynamics of nonlinear voter models.

We study the ordering dynamics of nonlinear voter models with multiple states, also providing a discussion of the two-state model. The rate with which an individual adopts an opinion scales as the qth power of the number of the individual's neighbors in that state. For q>1 the dynamics favor the opinion held by the most agents. The ordering to consensus is driven by deterministic drift, and noise plays only a minor role. For q<1 the dynamics favors minority opinions, and for multistate models the ordering proceeds through a noise-driven succession of metastable states. Unlike linear multistate systems, the nonlinear model cannot be reduced to an effective two-state model. We find that the average density of active interfaces in the model with multiple opinion states does not show a single exponential decay in time for q<1, again at variance with the linear model. This highlights the special character of the conventional (linear) voter model, in which deterministic drift is absent. As part of our analysis, we develop a pair approximation for the multistate model on graphs, valid for any positive real value of q, improving on previous approximations for nonlinear two-state voter models.

Noisy voter models in switching environments.

We study the stationary states of variants of the noisy voter model, subject to fluctuating parameters or external environments. Specifically, we consider scenarios in which the herding-to-noise ratio switches randomly and on different timescales between two values. We show that this can lead to a phase in which polarized and heterogeneous states exist. Second, we analyze a population of noisy voters subject to groups of external influencers, and show how multipeak stationary distributions emerge. Our work is based on a combination of individual-based simulations, analytical approximations in terms of a piecewise-deterministic Markov processes (PDMP), and on corrections to this process capturing intrinsic stochasticity in the linear-noise approximation. We also propose a numerical scheme to obtain the stationary distribution of PDMPs with three environmental states and linear velocity fields.

Niche overlap and Hopfield-like interactions in generalized random Lotka-Volterra systems.

We study communities emerging from generalized random Lotka-Volterra dynamics with a large number of species with interactions determined by the degree of niche overlap. Each species is endowed with a number of traits, and competition between pairs of species increases with their similarity in trait space. This leads to a model with random Hopfield-like interactions. We use tools from the theory of disordered systems, notably dynamic mean-field theory, to characterize the statistics of the resulting communities at stable fixed points and determine analytically when stability breaks down. Two distinct types of transition are identified in this way, both marked by diverging abundances but differing in the behavior of the integrated response function. At fixed points only a fraction of the initial pool of species survives. We numerically study the eigenvalue spectra of the interaction matrix between extant species. We find evidence that the two types of dynamical transition are, respectively, associated with the bulk spectrum or an outlier eigenvalue crossing into the right half of the complex plane.

Mutators can drive the evolution of multi-resistance to antibiotics.

Antibiotic combination therapies are an approach used to counter the evolution of resistance; their purported benefit is they can stop the successive emergence of independent resistance mutations in the same genome. Here, we show that bacterial populations with 'mutators', organisms with defects in DNA repair, readily evolve resistance to combination antibiotic treatment when there is a delay in reaching inhibitory concentrations of antibiotic-under conditions where purely wild-type populations cannot. In populations of Escherichia coli subjected to combination treatment, we detected a diverse array of acquired mutations, including multiple alleles in the canonical targets of resistance for the two drugs, as well as mutations in multi-drug efflux pumps and genes involved in DNA replication and repair. Unexpectedly, mutators not only allowed multi-resistance to evolve under combination treatment where it was favoured, but also under single-drug treatments. Using simulations, we show that the increase in mutation rate of the two canonical resistance targets is sufficient to permit multi-resistance evolution in both single-drug and combination treatments. Under both conditions, the mutator allele swept to fixation through hitch-hiking with single-drug resistance, enabling subsequent resistance mutations to emerge. Ultimately, our results suggest that mutators may hinder the utility of combination therapy when mutators are present. Additionally, by raising the rates of genetic mutation, selection for multi-resistance may have the unwanted side-effect of increasing the potential to evolve resistance to future antibiotic treatments.

Lyapunov exponents for temporal networks.

By interpreting a temporal network as a trajectory of a latent graph dynamical system, we introduce the concept of dynamical instability of a temporal network and construct a measure to estimate the network maximum Lyapunov exponent (nMLE) of a temporal network trajectory. Extending conventional algorithmic methods from nonlinear time-series analysis to networks, we show how to quantify sensitive dependence on initial conditions and estimate the nMLE directly from a single network trajectory. We validate our method for a range of synthetic generative network models displaying low- and high-dimensional chaos and finally discuss potential applications.

Breakdown of Random-Matrix Universality in Persistent Lotka-Volterra Communities.

The eigenvalue spectrum of a random matrix often only depends on the first and second moments of its elements, but not on the specific distribution from which they are drawn. The validity of this universality principle is often assumed without proof in applications. In this Letter, we offer a pertinent counterexample in the context of the generalized Lotka-Volterra equations. Using dynamic mean-field theory, we derive the statistics of the interactions between species in an evolved ecological community. We then show that the full statistics of these interactions, beyond those of a Gaussian ensemble, are required to correctly predict the eigenvalue spectrum and therefore stability. Consequently, the universality principle fails in this system. We thus show that the eigenvalue spectra of random matrices can be used to deduce the stability of "feasible" ecological communities, but only if the emergent non-Gaussian statistics of the interactions between species are taken into account.

Generalized Lotka-Volterra model with hierarchical interactions.

In the analysis of complex ecosystems it is common to use random interaction coefficients, which are often assumed to be such that all species are statistically equivalent. In this work we relax this assumption by imposing hierarchical interspecies interactions. These are incorporated into a generalized Lotka-Volterra dynamical system. In a hierarchical community species benefit more, on average, from interactions with species further below them in the hierarchy than from interactions with those above. Using dynamic mean-field theory, we demonstrate that a strong hierarchical structure is stabilizing, but that it reduces the number of species in the surviving community, as well as their abundances. Additionally, we show that increased heterogeneity in the variances of the interaction coefficients across positions in the hierarchy is destabilizing. We also comment on the structure of the surviving community and demonstrate that the abundance and probability of survival of a species are dependent on its position in the hierarchy.

Liquid-hexatic-solid phases in active and passive Brownian particles determined by stochastic birth and death events.

We study the effects of stochastic birth and death processes on the structural phases of systems of active and passive Brownian particles subject to volume exclusion. The total number of particles in the system is a fluctuating quantity, determined by the birth and death parameters and on the activity of the particles. As the birth and death parameters are varied, we find liquid, hexatic, and solid phases. For passive particles, these phases are found to be spatially homogeneous. For active particles, motility-induced phase separation (coexisting hexatic and liquid phases) occurs for large activity and sufficiently small birth rates. We also observe a reentrant transition to the hexatic phase when the birth rate is increased. This results from a balance of an increasing number of particles filling the system, and a larger number of defects resulting from the birth and death dynamics.

Local and global ordering dynamics in multistate voter models.

We investigate the time evolution of the density of active links and of the entropy of the distribution of agents among opinions in multistate voter models with all-to-all interaction and on uncorrelated networks. Individual realizations undergo a sequence of eliminations of opinions until consensus is reached. After each elimination the population remains in a metastable state. The density of active links and the entropy in these states varies from realization to realization. Making some simple assumptions we are able to analytically calculate the average density of active links and the average entropy in each of these states. We also show that, averaged over realizations, the density of active links decays exponentially, with a timescale set by the size and geometry of the graph, but independent of the initial number of opinion states. The decay of the average entropy is exponential only at long times when there are at most two opinions left in the population. Finally, we show how metastable states comprising only a subset of opinions can be artificially engineered by introducing precisely one zealot in each of the prevailing opinions.

Sampling rare trajectories using stochastic bridges.

The numerical quantification of the statistics of rare events in stochastic processes is a challenging computational problem. We present a sampling method that constructs an ensemble of stochastic trajectories that are constrained to have fixed start and end points (so-called stochastic bridges). We then show that by carefully choosing a set of such bridges and assigning an appropriate statistical weight to each bridge, one can focus more processing power on the rare events of a target stochastic process while faithfully preserving the statistics of these rare trajectories. Further, we also compare the stochastic bridges we produce to the Wentzel-Kramers-Brillouin (WKB) optimal paths of the target process, derived in the limit of low noise. We see that the generated paths, encoding the full statistics of the process, collapse onto the WKB optimal path as the level of noise is reduced. We propose that the method can also be used to judge the accuracy of the WKB approximation at finite levels of noise.

Eigenvalues of Random Matrices with Generalized Correlations: A Path Integral Approach.

Random matrix theory allows one to deduce the eigenvalue spectrum of a large matrix given only statistical information about its elements. Such results provide insight into what factors contribute to the stability of complex dynamical systems. In this Letter, we study the eigenvalue spectrum of an ensemble of random matrices with correlations between any pair of elements. To this end, we introduce an analytical method that maps the resolvent of the random matrix onto the response functions of a linear dynamical system. The response functions are then evaluated using a path integral formalism, enabling us to make deductions about the eigenvalue spectrum. Our central result is a simple, closed-form expression for the leading eigenvalue of a large random matrix with generalized correlations. This formula demonstrates that correlations between matrix elements that are not diagonally opposite, which are often neglected, can have a significant impact on stability.

Network meta-analysis and random walks.

Network meta-analysis (NMA) is a central tool for evidence synthesis in clinical research. The results of an NMA depend critically on the quality of evidence being pooled. In assessing the validity of an NMA, it is therefore important to know the proportion contributions of each direct treatment comparison to each network treatment effect. The construction of proportion contributions is based on the observation that each row of the hat matrix represents a so-called "evidence flow network" for each treatment comparison. However, the existing algorithm used to calculate these values is associated with ambiguity according to the selection of paths. In this article, we present a novel analogy between NMA and random walks. We use this analogy to derive closed-form expressions for the proportion contributions. A random walk on a graph is a stochastic process that describes a succession of random "hops" between vertices which are connected by an edge. The weight of an edge relates to the probability that the walker moves along that edge. We use the graph representation of NMA to construct the transition matrix for a random walk on the network of evidence. We show that the net number of times a walker crosses each edge of the network is related to the evidence flow network. By then defining a random walk on the directed evidence flow network, we derive analytically the matrix of proportion contributions. The random-walk approach has none of the associated ambiguity of the existing algorithm.

Analytical and Numerical Treatment of Continuous Ageing in the Voter Model.

The conventional voter model is modified so that an agent's switching rate depends on the 'age' of the agent-that is, the time since the agent last switched opinion. In contrast to previous work, age is continuous in the present model. We show how the resulting individual-based system with non-Markovian dynamics and concentration-dependent rates can be handled both computationally and analytically. The thinning algorithm of Lewis and Shedler can be modified in order to provide an efficient simulation method. Analytically, we demonstrate how the asymptotic approach to an absorbing state (consensus) can be deduced. We discuss three special cases of the age-dependent switching rate: one in which the concentration of voters can be approximated by a fractional differential equation, another for which the approach to consensus is exponential in time, and a third case in which the system reaches a frozen state instead of consensus. Finally, we include the effects of a spontaneous change of opinion, i.e., we study a noisy voter model with continuous ageing. We demonstrate that this can give rise to a continuous transition between coexistence and consensus phases. We also show how the stationary probability distribution can be approximated, despite the fact that the system cannot be described by a conventional master equation.

Beyond the adiabatic limit in systems with fast environments: A τ-leaping algorithm.

We propose a τ-leaping simulation algorithm for stochastic systems subject to fast environmental changes. Similar to conventional τ-leaping the algorithm proceeds in discrete time steps, but as a principal addition it captures environmental noise beyond the adiabatic limit. The key idea is to treat the input rates for the τ-leaping as (clipped) Gaussian random variables with first and second moments constructed from the environmental process. In this way, each step of the algorithm retains environmental stochasticity to subleading order in the timescale separation between system and environment. We test the algorithm on several toy examples with discrete and continuous environmental states and find good performance in the regime of fast environmental dynamics. At the same time, the algorithm requires significantly less computing time than full simulations of the combined system and environment. In this context we also discuss several methods for the simulation of stochastic population dynamics in time-varying environments with continuous states.

Switching environments, synchronous sex, and the evolution of mating types.

While facultative sex is common in sexually reproducing species, for reasons of tractability most mathematical models assume that such sex is asynchronous in the population. In this paper, we develop a model of switching environments to instead capture the effect of an entire population transitioning synchronously between sexual and asexual modes of reproduction. We use this model to investigate the evolution of the number of self-incompatible mating types in finite populations, which empirically can range from two to thousands. When environmental switching is fast, we recover the results of earlier studies that implicitly assumed populations were engaged in asynchronous sexual reproduction. However when the environment switches slowly, we see deviations from previous asynchronous theory, including a lower number of mating types at equilibrium and bimodality in the stationary distribution of mating types. We provide analytic approximations for both the fast and slow switching regimes, as well as a numerical scheme based on the Kolmogorov equations for the system to quickly evaluate the model dynamics at intermediate parameters. Our approach exploits properties of integer partitions in number theory. We also demonstrate how additional biological processes such as selective sweeps can be accounted for in this switching environment framework, showing that beneficial mutations can further erode mating type diversity in synchronous facultatively sexual populations.

Geospatial distributions reflect temperatures of linguistic features.

Quantifying the speed of linguistic change is challenging because the historical evolution of languages is sparsely documented. Consequently, traditional methods rely on phylogenetic reconstruction. Here, we propose a model-based approach to the problem through the analysis of language change as a stochastic process combining vertical descent, spatial interactions, and mutations in both dimensions. A notion of linguistic temperature emerges naturally from this analysis as a dimensionless measure of the propensity of a linguistic feature to undergo change. We demonstrate how temperatures of linguistic features can be inferred from their present-day geospatial distributions, without recourse to information about their phylogenies. Thus, the evolutionary dynamics of language, operating across thousands of years, leave a measurable geospatial signature. This signature licenses inferences about the historical evolution of languages even in the absence of longitudinal data.

Zealots in multistate noisy voter models.

The noisy voter model is a stylized representation of opinion dynamics. Individuals copy opinions from other individuals, and are subject to spontaneous state changes. In the case of two opinion states this model is known to have a noise-driven transition between a unimodal phase, in which both opinions are present, and a bimodal phase, in which one of the opinions dominates. The presence of zealots can remove the unimodal and bimodal phases in the model with two opinion states. Here we study the effects of zealots in noisy voter models with M>2 opinion states on complete interaction graphs. We find that the phase behavior diversifies, with up to six possible qualitatively different types of stationary states. The presence of zealots removes some of these phases, but not all. We analyze situations in which zealots affect the entire population, or only a fraction of agents, and show that this situation corresponds to a single-community model with a fractional number of zealots, further enriching the phase diagram. Our study is conducted analytically based on effective birth-death dynamics for the number of individuals holding a given opinion. Results are confirmed in numerical simulations.

Degree irregularity and rank probability bias in network meta-analysis.

Network meta-analysis (NMA) is a statistical technique for the comparison of treatment options. Outcomes of Bayesian NMA include estimates of treatment effects, and the probabilities that each treatment is ranked best, second best and so on. How exactly network topology affects the accuracy and precision of these outcomes is not fully understood. Here we carry out a simulation study and find that disparity in the number of trials involving different treatments leads to a systematic bias in estimated rank probabilities. This bias is associated with an increased variation in the precision of treatment effect estimates. Using ideas from the theory of complex networks, we define a measure of "degree irregularity" to quantify asymmetry in the number of studies involving each treatment. Our simulations indicate that more regular networks have more precise treatment effect estimates and smaller bias of rank probabilities. Conversely, these topological effects are not observed for the accuracy of treatment effect estimates. This reinforces the importance of taking into account multiple measures, rather than making decisions based on a single metric. We also find that degree regularity is a better indicator for the accuracy and precision of parameter estimates in NMA than both the total number of studies in a network and the disparity in the number of trials per comparison. These results have implications for planning future trials. We demonstrate that choosing trials which reduce the network's irregularity can improve the precision and accuracy of parameter estimates from NMA.

Competition delays multi-drug resistance evolution during combination therapy.

Combination therapies have shown remarkable success in preventing the evolution of resistance to multiple drugs, including HIV, tuberculosis, and cancer. Nevertheless, the rise in drug resistance still remains an important challenge. The capability to accurately predict the emergence of resistance, either to one or multiple drugs, may help to improve treatment options. Existing theoretical approaches often focus on exponential growth laws, which may not be realistic when scarce resources and competition limit growth. In this work, we study the emergence of single and double drug resistance in a model of combination therapy of two drugs. The model describes a sensitive strain, two types of single-resistant strains, and a double-resistant strain. We compare the probability that resistance emerges for three growth laws: exponential growth, logistic growth without competition between strains, and logistic growth with competition between strains. Using mathematical estimates and numerical simulations, we show that between-strain competition only affects the emergence of single resistance when resources are scarce. In contrast, the probability of double resistance is affected by between-strain competition over a wider space of resource availability. This indicates that competition between different resistant strains may be pertinent to identifying strategies for suppressing drug resistance, and that exponential models may overestimate the emergence of resistance to multiple drugs. A by-product of our work is an efficient strategy to evaluate probabilities of single and double resistance in models with multiple sequential mutations. This may be useful for a range of other problems in which the probability of resistance is of interest.

Dispersal-induced instability in complex ecosystems.

In his seminal work in the 1970s, Robert May suggested that there is an upper limit to the number of species that can be sustained in stable equilibrium by an ecosystem. This deduction was at odds with both intuition and the observed complexity of many natural ecosystems. The so-called stability-diversity debate ensued, and the discussion about the factors contributing to ecosystem stability or instability continues to this day. We show in this work that dispersal can be a destabilising influence. To do this, we combine ideas from Alan Turing's work on pattern formation with May's random-matrix approach. We demonstrate how a stable equilibrium in a complex ecosystem with trophic structure can become unstable with the introduction of dispersal in space, and we discuss the factors which contribute to this effect. Our work highlights that adding more details to the model of May can give rise to more ways for an ecosystem to become unstable. Making May's simple model more realistic is therefore unlikely to entirely remove the upper bound on complexity.