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Current epidemics in the biological and social domains, such as the COVID-19 pandemic or the spread of misinformation, are challenging many of the classic assumptions of mathematical contagion models. Chief among them are the complex patterns of transmission caused by heterogeneous group sizes and infection risk varying by orders of magnitude in different settings, like indoor versus outdoor gatherings in the COVID-19 pandemic or different moderation practices in social media communities. Here, we include these novel features in an epidemic model on a weighted hypergraph to capture group-specific transmission rates. We find an induced superlinear force of infection during the emergence of a new outbreak. The dynamics produced at the individual and group levels are therefore more similar to complex contagions than the simple contagions of classic disease models. This result is an important cautionary tale for the challenging task of inferring transmission mechanisms from incidence data. Yet, it also opens the door to effective models that capture complex features of epidemics through nonlinear forces of infection., 16 pages, 4 figures

Starting from a stochastic individual based description of an SIS epidemic spreading on a random network, we study the dynamics when the size of the network tends to infinity. We recover in the limit an infinite-dimensional integro-differential equation studied by Delmas, Dronnier and Zitt (2022) for an SIS epidemic propagating on a graphon. Our work covers the case of dense and sparse graphs, when the number of edges is of order $\Theta(n^a)$ with $a>1$, but not the case of very sparse graphs with $a=1$. In order to establish our limit theorem, we have to deal with both the convergence of the graph to the graphon and the convergence of the stochastic process spreading on top of these random structures: in particular, we propose a coupling between the process of interest and an epidemic that spreads on the complete graph but with a modified infectivity., Comment: Keywords: random graph, mathematical model for epidemiology, measure-valued process, large network limit, limit theorem, dense graph, sparse graph. Acknowledgments: This work was financed by the Labex B\'ezout (ANR-10-LABX-58) and the platform MODCOV19 of the National Institute of Mathematical Sciences and their Interactions of CNRS. 30 pages, including an appendix of 5 pages

Previous investigations into creative and innovation networks have suggested that innovations often occurs at the boundary between the network's core and periphery. In this work, we investigate the effect of global core-periphery network structure on the speed and quality of cultural innovation. Drawing on differing notions of core-periphery structure from [arXiv:1808.07801] and [doi:10.1016/S0378-8733(99)00019-2], we distinguish decentralized core-periphery, centralized core-periphery, and affinity network structure. We generate networks of these three classes from stochastic block models (SBMs), and use them to run an agent-based model (ABM) of collective cultural innovation, in which agents can only directly interact with their network neighbors. In order to discover the highest-scoring innovation, agents must discover and combine the highest innovations from two completely parallel technology trees. We find that decentralized core-periphery networks outperform the others by finding the final crossover innovation more quickly on average. We hypothesize that decentralized core-periphery network structure accelerates collective problem-solving by shielding peripheral nodes from the local optima known by the core community at any given time. We then build upon the "Two Truths" hypothesis regarding community structure in spectral graph embeddings, first articulated in [arXiv:1808.07801], which suggests that the adjacency spectral embedding (ASE) captures core-periphery structure, while the Laplacian spectral embedding (LSE) captures affinity. We find that, for core-periphery networks, ASE-based resampling best recreates networks with similar performance on the innovation SBM, compared to LSE-based resampling. Since the Two Truths hypothesis suggests that ASE captures core-periphery structure, this result further supports our hypothesis., 9 pages, 5 figures, AAMAS 2023 (accepted)

The genetic code refers to a rule that maps 64 codons to 20 amino acids. Nearly all organisms, with few exceptions, share the same genetic code, the standard genetic code (SGC). While it remains unclear why this universal code has arisen and been maintained during evolution, it may have been preserved under selection pressure. Theoretical studies comparing the SGC and numerically created hypothetical random genetic codes have suggested that the SGC has been subject to strong selection pressure for being robust against translation errors. However, these prior studies have searched for random genetic codes in only a small subspace of the possible code space due to limitations in computation time. Thus, how the genetic code has evolved, and the characteristics of the genetic code fitness landscape, remain unclear. By applying multicanonical Monte Carlo, an efficient rare-event sampling method, we efficiently sampled random codes from a much broader random ensemble of genetic codes than in previous studies, estimating that only one out of every $10^{20}$ random codes is more robust than the SGC. This estimate is significantly smaller than the previous estimate, one in a million. We also characterized the fitness landscape of the genetic code that has four major fitness peaks, one of which includes the SGC. Furthermore, genetic algorithm analysis revealed that evolution under such a multi-peaked fitness landscape could be strongly biased toward a narrow peak, in an evolutionary path-dependent manner., Comment: 8 pages, 3 figures

In their well-known textbook (Vincent & Brown, 2005), Vincent and Brown suggested an attractive approach for studying evolutionary dynamics of populations that are heterogeneous with respect to some strategy that affects the fitness of individuals in the population. The authors developed a theory, whose goal was to expand the applicability of mathematical models of population dynamics by including dynamics of an evolving heritable phenotype trait subject to natural selection. The authors studied both the case of evolution of individual traits and of mean traits in the population (or species) and the dynamics of total population size. The authors consider the developed approach as (more or less) universally applicable to models with any fitness function and any initial distribution of strategies, which is symmetric and has small variance. Here it was shown that the scope of the approach proposed by Vincent & Brown is unfortunately much more limited. I show that the approach gives exact results only if the population dynamics linearly depends on the trait; examples where the approach is incorrect are given., 39 pages, 11 figures

Many reaction-diffusion models produce travelling wave solutions that can be interpreted as waves of invasion in biological scenarios such as wound healing or tumour growth. These partial differential equation models have since been adapted to describe the interactions between cells and extracellular matrix (ECM), using a variety of different underlying assumptions. In this work, we derive a system of reaction-diffusion equations, with cross-species density-dependent diffusion, by coarse-graining an agent-based, volume-filling model of cell invasion into ECM. We study the resulting travelling wave solutions both numerically and analytically across various parameter regimes. Subsequently, we perform a systematic comparison between the behaviours observed in this model and those predicted by simpler models in the literature that do not take into account volume-filling effects in the same way. Our study justifies the use of some of these simpler, more analytically tractable models in reproducing the qualitative properties of the solutions in some parameter regimes, but it also reveals some interesting properties arising from the introduction of cell and ECM volume-filling effects, where standard model simplifications might not be appropriate., 28 pages, 12 figures

Genetic algorithm (GA) is inspired by biological evolution of genetic organisms by optimizing the genotypic combinations encoded within each individual with the help of evolutionary operators, suggesting that GA may be a suitable model for studying real-life evolutionary processes. This paper describes the design of a Python library for artificial life simulation, Digital Organism Simulation Environment (DOSE), based on GA and biological hierarchy starting from genetic sequence to population. A 3-character instruction set that does not take any operand is introduced as genetic code for digital organism. This mimics the 3-nucleotide codon structure in naturally occurring DNA. In addition, the context of a 3-dimensional world composing of ecological cells is introduced to simulate a physical ecosystem. Using DOSE, an experiment to examine the changes in genetic sequences with respect to mutation rates is presented.

A physics-informed neural network (PINN) embedded with the susceptible-infected-removed (SIR) model is devised to understand the temporal evolution dynamics of infectious diseases. Firstly, the effectiveness of this approach is demonstrated on synthetic data as generated from the numerical solution of the susceptible-asymptomatic-infected-recovered-dead (SAIRD) model. Then, the method is applied to COVID-19 data reported for Germany and shows that it can accurately identify and predict virus spread trends. The results indicate that an incomplete physics-informed model can approach more complicated dynamics efficiently. Thus, the present work demonstrates the high potential of using machine learning methods, e.g., PINNs, to study and predict epidemic dynamics in combination with compartmental models., 23 pages, 14 figures

In some patients of myeloproliferative neoplasm, two genetic mutations can be found: JAK2 V617F and TET2. When one mutation is present or not, the other mutation has different effects on regulating gene expressions. Besides, when both mutations are present, the order of occurrence might make a difference. In this paper, we build nonlinear ordinary differential equation models and Markov chain models to explain such phenomena., Highly incomplete work that needs extensive revision

In a Bienaym\'{e}-Galton-Watson process for which there is a positiveprobability for individuals of having no offspring, there is a subtlebalance and dependence between the sterile nodes (the dead nodes or leaves)and the prolific ones (the productive nodes) both at and up to the currentgeneration. We explore the many facets of this problem, especially in thecontext of an exactly solvable linear-fractional branching mechanism at allgeneration. Eased asymptotic issues are investigated. Relation of thisspecial branching process to skip-free to the left and simple random walks'excursions is then investigated. Mutual statistical information on theirshapes can be learnt from this association.

We introduce a multi-allele Wright-Fisher model with non-recurrent, reversible mutation and directional selection. In this setting, the allele frequencies at a single locus track the path of a hybrid jump-diffusion process with state space given by the vertex and edge set of a graph. Vertices represent monomorphic population states and edge-positions mark the biallelic proportions of ancestral and derived alleles during polymorphic segments. We derive the stationary distribution in mutation-selection-drift equilibrium and obtain the expected allele frequency spectrum under large population size scaling. For the extended model with multiple independent loci we derive rigorous upper bounds for a wide class of associated measures of genetic variation. Within this framework we present mathematically precise arguments to conclude that the presence of directional selection reduces the magnitude of genetic variation, as constrained by the bounds for neutral evolution.

A long-standing goal of evolutionary biology is to decode how gene regulatory processes contribute to organismal diversity, both within and between species. This question has remained challenging to answer, due both to the difficulties of predicting function from non-coding sequence, and to the technological constraints of laboratory research with non-model taxa. However, a recent methodological development in functional genomics, the massively parallel reporter assay (MPRA), makes it possible to test thousands to millions of sequences for regulatory activity in a single in vitro experiment. It does so by combining traditional, single-locus episomal reporter assays (e.g., luciferase reporter assays) with the scalability of high-throughput sequencing. In this perspective, we discuss the execution, advantages, and limitations of MPRAs for research in evolutionary biology. We review recent studies that have made use of this approach to address explicitly evolutionary questions, highlighting study designs that we believe are particularly well-positioned to gain from MPRA approaches. Additionally, we propose solutions for extending these powerful assays to rare taxa and those with limited genomic resources. In doing so, we underscore the broad potential of MPRAs to drive genome-scale functional evolutionary genetics studies in non-traditional model organisms.

The bacterium Xylella fastidiosa (Xf) is mainly transmitted by the spittlebug, Philaenus spumarius, in Europe, where it has caused significant economic damage to olive and almond trees. Understanding the factors that determine disease dynamics in pathosystems that share similarities can help design control strategies focused on minimizing transmission chains. Here we introduce a compartmental model for Xf-caused diseases in Europe that accounts for the main relevant epidemiological processes, including the seasonal dynamics of P. spumarius. The model was confronted with epidemiological data from the two major outbreaks of Xf in Europe, the olive quick disease syndrome (OQDS) in Apulia, Italy, caused by the subspecies pauca, and the almond leaf scorch disease (ALSD) in Majorca, Spain, caused by subspecies multiplex and fastidiosa. Using a Bayesian inference framework, we show how the model successfully reproduces the general field data in both diseases. In a global sensitivity analysis, the vector-plant and plant-vector transmission rates, together with the vector removal rate, were the most influential parameters in determining the time of the infected host population peak, the incidence peak and the final number of dead hosts. We also used our model to check different vector-based control strategies, showing that a joint strategy focused on increasing the rate of vector removal while lowering the number of annual newborn vectors is optimal for disease control., 18 pages, 8 figures

A reaction-diffusion Leslie-Gower predator-prey model, incorporating the fear effect and prey refuge, with Beddington-DeAngelis functional response, is introduced. A qualitative analysis of the solutions of the model and the stability analysis of the coexistence equilibrium, are performed. Sufficient conditions guaranteeing the occurrence of Turing instability have been determined either in the case of self-diffusion or in the case of cross-diffusion. Different types of Turing patterns, representing a spatial redistribution of population in the environment, emerge for different values of the model parameters.

According to the evolutionary death-birth protocol, a player is chosen randomly to die and neighbors compete for the available position proportional to their fitness. Hence, the status of the focal player is completely ignored and has no impact on the strategy update. In this work, we revisit and generalize this rule by introducing a weight factor to compare the payoff values of the focal and invading neighbors. By means of evolutionary graph theory, we analyze the model on joint transitive graphs to explore the possible consequences of the presence of a weight factor. We find that focal weight always hinders cooperation under weak selection strength. Surprisingly, the results show a non-trivial tipping point of the weight factor where the threshold of cooperation success shifts from positive to negative infinity. Once focal weight exceeds this tipping point, cooperation becomes unreachable. Our theoretical predictions are confirmed by Monte Carlo simulations on a square lattice of different sizes. We also verify the robustness of the conclusions to arbitrary two-player prisoner's dilemmas, to dispersal graphs with arbitrary edge weights, and to interaction and dispersal graphs overlapping arbitrarily., Comment: 8 two-column pages, 2 figures, accepted for publication in Physical Review E

Combining a spatiotemporal, multi-agent based model of a foraging ecosystem with linear, genetically programmed rules for the agents' behaviors results in implicit, endogenous, objective functions and selection algorithms based on "natural selection". Use of this implicit optimization of genetic programs for study of biological systems is tested by application to an artificial foraging ecosystem, and compared with established biological, ecological, and stochastic gene diffusion models. Limited program memory and execution time constraints emulate real-time and concurrent properties of physical and biological systems, and stress test the optimization algorithms. Relative fitness of the agents' programs and efficiency of the resultant populations as functions of these constraints gauge optimization effectiveness and efficiency. Novel solutions confirm the creativity of the optimization process and provide an unique opportunity to experimentally test the neutral code bloating hypotheses. Use of this implicit, endogenous, evolutionary optimization of spatially interacting, genetically programmed agents is thus shown to be novel, consistent with biological systems, and effective and efficient in discovering fit and novel solutions., 1 table, 4 figures, 1 PDL

Researchers have employed stochastic simulations to determine the validity of their theoretical findings and to study analytically intractable spreading dynamics. In both cases, the correctness and efficiency of the simulation algorithm are of paramount importance. We prove in this article that the Next Reaction Method and the non-Markovian Gillespie algorithm, two algorithms for simulating non-Markovian epidemics, are statistically equivalent. We also study the performance and applicability under various circumstances through complexity analyses and numerical experiments. In our numerical simulations, we apply the Next Reaction Method and the Gillespie algorithm to epidemic simulations on time-varying networks and epidemic simulations with cooperative infections. Both tasks have only been done using the Gillespie algorithm, while we show that the Next Reaction Method is a good alternative. We believe this article may also serve as a guide for choosing simulation algorithms that are both correct and efficient for researchers from epidemiology and beyond.

The advances in variational inference are providing promising paths in Bayesian estimation problems. These advances make variational phylogenetic inference an alternative approach to Markov Chain Monte Carlo methods for approximating the phylogenetic posterior. However, one of the main drawbacks of such approaches is the modelling of the prior through fixed distributions, which could bias the posterior approximation if they are distant from the current data distribution. In this paper, we propose an approach and an implementation framework to relax the rigidity of the prior densities by learning their parameters using a gradient-based method and a neural network-based parameterization. We applied this approach for branch lengths and evolutionary parameters estimation under several Markov chain substitution models. The results of performed simulations show that the approach is powerful in estimating branch lengths and evolutionary model parameters. They also show that a flexible prior model could provide better results than a predefined prior model. Finally, the results highlight that using neural networks improves the initialization of the optimization of the prior density parameters., Accepted as a full paper for publication at RECOMB-CG 2023 (Camera-ready version). 15 pages (excluding references), 6 tables and 1 figure

Epidemic propagation on networks represents an important departure from traditional massaction models. However, the high-dimensionality of the exact models poses a challenge to both mathematical analysis and parameter inference. By using mean-field models, such as the pairwise model (PWM), the complexity becomes tractable. While such models have been used extensively for model analysis, there is limited work in the context of statistical inference. In this paper, we explore the extent to which the PWM with the susceptible-infected-recovered (SIR) epidemic can be used to infer disease- and network-related parameters. The widely-used MLE approach exhibits several issues pertaining to parameter unidentifiability and a lack of robustness to exact knowledge about key quantities such as population size and/or proportion of under reporting. As an alternative, we considered the recently developed dynamical survival analysis (DSA). For scenarios in which there is no model mismatch, such as when data are generated via simulations, both methods perform well despite strong dependence between parameters. However, for real-world data, such as foot-and-mouth, H1N1 and COVID19, the DSA method appears more robust to potential model mismatch and the parameter estimates appear more epidemiologically plausible. Taken together, however, our findings suggest that network-based mean-field models can be used to formulate approximate likelihoods which, coupled with an efficient inference scheme, make it possible to not only learn about the parameters of the disease dynamics but also that of the underlying network., 23 pages, 15 figures

Dengue and Zika incidence data and the latest research have raised questions about how dengue vaccine strategies might be impacted by the emergence of Zika virus. Existing antibodies to one virus might temporarily protect or promote infection by the other through antibody-dependent enhancement (ADE). With this condition, understanding the dynamics of propagation of these two viruses is of great importance when implementing vaccines. In this work, we analyze the effect of vaccination against one strain, in a two-strain model that accounts for cross-immunity and ADE. Using basic and invasion reproductive numbers, we examined the dynamics of the model and provide conditions to ensure the stability of the disease-free equilibrium. We provide conditions on cross-immunity, ADE and vaccination rate under which the vaccination could ensure the global stability of the disease-free equilibrium. The results indicate scenarios in which vaccination against one strain may improve or worsen the control of the other, as well as contribute to the eradication or persistence of one or both viruses in the population., Corrected typos. Revised figures, results unchanged

In this paper, we discuss a hierarchical model, based on a Ricker competition model. The species considered are competing for resources and may be subject to an Allee effect due to mate limitation, anti-predator vigilance or aggression, cooperative predation or resource defense, or social thermoregulation. The species may be classified into a more dominant species and less dominant or ``wimpy" species or just as a predator and prey. The model under consideration also has components taking into account immigration in both species and a more structured Holling type II functional response. Local and global stability analyses are discussed and simulations are provided. We also consider demographic stochasticity on the species due to environmental fluctuations in the form of Wiener processes and we show that there are conditions under which a global solution exists, stationary distributions exists, and strong persistence in mean of the species is possible. We also use Wasserstein distance to show empirically that stochasticity can acts as bifurcation parameter.

In biology, stochastic branching processes with a two-stage, hierarchical structure arise in the study of population dynamics, gene expression, and phylogenetic inference. These models have been commonly analyzed using generating functions, the method of characteristics and various perturbative approximations. Here we describe a general method for analyzing hierarchic first-order reaction networks using Lie theory. Crucially, we identify the fact that the Lie group associated to hierarchic reaction networks decomposes as a wreath product of the groups associated to the subnetworks of the independent and dependent types. After explaining the general method, we illustrate it on a model of population dynamics and the so-called two-state or telegraph model of single-gene transcription. Solutions to such processes provide essential input to downstream methods designed to attempt to infer parameters of these and related models., 9 pages

Over 80% of human cancers originate from the epithelium, which covers the outer and inner surfaces of organs and blood vessels. In stratified epithelium, the bottom layers are occupied by stem and stem-like cells that continually divide and replenish the upper layers. In this work, we study the spread of premalignant mutant clones and cancer initiation in stratified epithelium using the biased voter model on stacked two-dimensional lattices. Our main result is an estimate of the propagation speed of a premalignant mutant clone, which is asymptotically precise in the cancer-relevant weak-selection limit. We use our main result to study cancer initiation under a two-step mutational model of cancer, which includes computing the distributions of the time of cancer initiation and the size of the premalignant clone giving rise to cancer. Our work quantifies the effect of epithelial tissue thickness on the process of carcinogenesis, thereby contributing to an emerging understanding of the spatial evolutionary dynamics of cancer., Comment: 49 pages, 11 figures, accepted for publication in Annals of Applied Probability

The rooted subtree prune and regraft (rSPR) distance between two rooted binary phylogenetic trees is a well-studied measure of topological dissimilarity that is NP-hard to compute. Here we describe an improved linear kernel for the problem. In particular, we show that if the classical subtree and chain reduction rules are augmented with a modified type of chain reduction rule, the resulting trees have at most 9k−3 leaves, where k is the rSPR distance; and that this bound is tight. In comparison, the previous best-known linear kernel is of size at most 28k. To achieve this improvement we introduce cyclic generators, which can be viewed as cyclic analogues of the generators used in the phylogenetic networks literature. As a corollary to our main result we also give an improved weighted linear kernel for the minimum hybridization problem on two rooted binary phylogenetic trees.

The algebraic properties of flattenings and subflattenings provide direct methods for identifying edges in the true phylogeny -- and by extension the complete tree -- using pattern counts from a sequence alignment. The relatively small number of possible internal edges among a set of taxa (compared to the number of binary trees) makes these methods attractive, however more could be done to evaluate their effectiveness for inferring phylogenetic trees. This is the case particularly for subflattenings, and our work makes progress in this area. We introduce software for constructing and evaluating subflattenings for splits, utilising a number of methods to make computing subflattenings more tractable. We then present the results of simulations we have performed in order to compare the effectiveness of subflattenings to that of flattenings in terms of split score distributions, and susceptibility to possible biases. We find that subflattenings perform similarly to flattenings in terms of the distribution of split scores on the trees we examined, but may be less affected by bias arising from both split size/balance and long branch attraction. These insights are useful for developing effective algorithms to utilise these tools for the purpose of inferring phylogenetic trees., Comment: 21 pages, 13 figures

Predator-prey models have been shown to exhibit resonance-like behaviour, in which random fluctuations in the number of organisms (demographic noise) are amplified when their frequency is close to the natural oscillatory frequency of the system. This behaviour has been traditionally studied in models with exponentially distributed replication and death times. Here we consider a biologically more realistic model, in which organisms replicate quasi-synchronously such that the distribution of replication times has a narrow maximum at some $T>0$ corresponding to the mean doubling time. We show that when the frequency of replication $f=1/T$ is tuned to the natural oscillatory frequency of the predator-prey model, the system exhibits oscillations that are much stronger than in the model with Poissonian (non-synchronous) replication and death. The effect can be explained by resonant amplification of coloured noise generated by quasi-synchronous replication events. To show this, we consider a single-species model with quasi-synchronous replication. We calculate the spectrum and the amplitude of demographic noise in this model, and use these results to obtain these quantities for the two-species model., 14 pages, 12 figures

The soil seed bank is a major component of plant communities. However, long-term analyses of the dynamics of the seed bank and the ensuing vegetation are rare. Here, we studied the dynamics in plant communities with high dominance of annuals in Mediterranean, semiarid, and arid ecosystems for nine consecutive years. For annuals, we hypothesized that the density of the seed bank would be more stable than the density of the standing herbaceous vegetation. Moreover, we predicted that differences in temporal variability between the seed bank and the vegetation would increase with aridity, where year-to-year rainfall variability is higher. We found that the temporal variability at the population level (assessed as the standard deviation of the loge-transformed density) of the nine dominant annuals in each site did not differ between the seed bank and the ensuing vegetation in any of the sites. For the total density of annuals, patterns depended on aridity. In the Mediterranean site, the temporal variability was similar in the seed bank and the vegetation (0.40 vs. 0.40). Still, in the semiarid and arid sites, variability in the seed bank was lower than in the vegetation (0.49 vs. 1.01 and 0.63 vs. 1.38, respectively). This difference between the population-level patterns and the total density of annuals can be related to the lower population synchrony in their seed bank. In contrast, for the herbaceous perennials (all species combined), the seed bank variability was higher than in the vegetation. Overall, our results highlight the role of the seed bank in buffering the annual vegetation density with increasing climatic uncertainty typical in aridity gradients. This role is crucial under the increasing uncertainty imposed by climatic change in the region.

Bacteriophages are central to microbial ecosystems for balancing bacterial populations and promoting evolution by applying strong selection pressure. Here we review some of the known aspects that modulate phage-bacteria interaction in a way that naturally promotes their coexistence. We focus on the modulations that arise from structural, physical, or physiological constraints. We argue they should play roles in many phage-bacteria systems providing sustainable diversity., Short review. 7 pages, 3 figures. Minor update of the texts

We study communities emerging from generalised random Lotka--Volterra dynamics with a large number of species and with competitive exclusion. 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 Hebbian interactions. We use tools from the theory of disordered systems, notably dynamic mean field theory, to characterise 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 behaviour 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., 14 pages, 9 figures + Supplement

Phylodynamics is a set of population genetics tools that aim at reconstructing demographic history of a population based on molecular sequences of individuals sampled from the population of interest. One important task in phylodynamics is to estimate changes in (effective) population size. When applied to infectious disease sequences such estimation of population size trajectories can provide information about changes in the number of infections. To model changes in the number of infected individuals, current phylodynamic methods use non-parametric approaches, parametric approaches, and stochastic modeling in conjunction with likelihood-free Bayesian methods. The first class of methods yields results that are hard-to-interpret epidemiologically. The second class of methods provides estimates of important epidemiological parameters, such as infection and removal/recovery rates, but ignores variation in the dynamics of infectious disease spread. The third class of methods is the most advantageous statistically, but relies on computationally intensive particle filtering techniques that limits its applications. We propose a Bayesian model that combines phylodynamic inference and stochastic epidemic models, and achieves computational tractability by using a linear noise approximation (LNA) --- a technique that allows us to approximate probability densities of stochastic epidemic model trajectories. LNA opens the door for using modern Markov chain Monte Carlo tools to approximate the joint posterior distribution of the disease transmission parameters and of high dimensional vectors describing unobserved changes in the stochastic epidemic model compartment sizes (e.g., numbers of infectious and susceptible individuals). We apply our estimation technique to Ebola genealogies estimated using viral genetic data from the 2014 epidemic in Sierra Leone and Liberia., Comment: 43 pages, 6 figures in the main text

Livestock mobility, particularly that of small and large ruminants, is one of the main pillars of production and trade in West Africa: livestock is moved around in search of better grazing or sold in markets for domestic consumption and for festival-related activities. These movements cover several thousand kilometers and have the capability of connecting the whole West African region thus facilitating the diffusion of many animal and zoonotic diseases. Several factors shape mobility patterns even in normal years and surveillance systems need to account for such changes. In this paper, we present a procedure based on temporal network theory to identify possible sentinel locations using two indicators: vulnerability (i.e. the probability of being reached by the disease) and time of infection (i.e. the time of first arrival of the disease). Using these indicators in our structural analysis of the changing network enabled us to identify a set of nodes that could be used in an early warning system. As a case study we simulated the introduction of F.A.S.T. (Foot and Mouth Similar Transboundary) diseases in Senegal and used data taken from 2020 Sanitary certificates (LPS, laissez-passer sanitaire) issued by the Senegalese Veterinary Services to reconstruct the national mobility network. Our analysis showed that a static approach can significantly overestimate the speed and the extent of disease propagation, whereas temporal analysis revealed that the reachability and vulnerability of the different administrative departments (used as nodes of the mobility network) change over the course of the year. For this reason, several sets of sentinel nodes were identified in different periods of the year, underlining the role of temporality in shaping patterns of disease diffusion., 36 pages (25 main manuscript, 11 supplementary information), 17 figures (7 main manuscript, 10 supplementary information), submitted to Transboundary and Emerging Diseases

Non-standard Lagrangians do not display any discernible energy-like terms, yet they give the same equations of motion as standard Lagrangians, which have easily identifiable energy-like terms. A new method to derive non-standard Lagrangians for second-order nonlinear differential equations with damping is developed and the limitations of this method are explored. It is shown that the limitations do not exist only for those nonlinear dynamical systems that can be converted into linear ones. The obtained results are applied to selected population dynamics models for which non-standard Lagrangians and their corresponding null Lagrangians and gauge functions are derived, and their roles in the population dynamics are discussed., 16 pages, 2 tables

Discrete time, spatially extended models play an important role in ecology, modelling population dynamics of species ranging from micro-organisms to birds. An important question is how 'bottom up', individual-based models can be approximated by 'top down' models of dynamics. Here, we study a class of spatially explicit individual-based models with contest competition: where species compete for space in local cells and then disperse to nearby cells. We start by describing simulations of the model, which exhibit large-scale discrete oscillations and characterise these oscillations by measuring spatial correlations. We then develop two new approximate descriptions of the resulting spatial population dynamics. The first is based on local interactions of the individuals and allows us to give a difference equation approximation of the system over small dispersal distances. The second approximates the long-range interactions of the individual-based model. These approximations capture demographic stochasticity from the individual-based model and show that dispersal stabilizes population dynamics. We calculate extinction probability for the individual-based model and show convergence between the local approximation and the non-spatial global approximation of the individual-based model as dispersal distance and population size simultaneously tend to infinity. Our results provide new approximate analytical descriptions of a complex bottom-up model and deepen understanding of spatial population dynamics.

Since the foundation of population genetics, it is believed that directional selection should reduce genetic diversity. We present an exactly solvable population model which contradicts this intuition. The population is modelled as a cloud of particles evolving in a 1-dimensional fitness space (fitness wave). We show the existence of a phase transition which separates the parameter space into a weak and a strong selection regimes. We find that genetic diversity is highly non-monotone in the selection strength and, in contrast with the common intuition, our model predicts that genetic diversity is typically higher in the strong selection regime. This apparent paradox is resolved by observing that a higher selection strength increases the absolute fitness of the wave, but typically generate lower relative fitness between individuals within the wave. These findings entail that inferring the magnitude of natural selection from genetic data may raise some serious conceptual issues. Along the way, we uncover a new phase transition in front propagation. Namely, we show that the transition from weak to strong selection can be reformulated in terms of a transition from fully-pulled to semi-pulled waves. This transition is the pulled analog to the semi-pushed to fully-pushed regimes observed in noisy-FKPP travelling waves in the presence of Allee effect.

Large-scale events like the UEFA Euro~2020 football (soccer) championship offer a unique opportunity to quantify the impact of gatherings on the spread of COVID-19, as the number and dates of matches played by participating countries resembles a randomized study. Using Bayesian modeling and the gender imbalance in COVID-19 data, we attribute 840,000 (95% CI: [0.39M, 1.26M]) COVID-19 cases across 12 countries to the championship. The impact depends non-linearly on the initial incidence, the reproduction number $R$, and the number of matches played. The strongest effects are seen in Scotland and England, where as much as 10,000 primary cases per million inhabitants occur from championship-related gatherings. The average match-induced increase in $R$ was 0.46 [0.18, 0.75] on match days, but important matches caused an increase as large as +3. Altogether, our results provide quantitative insights that help judge and mitigate the impact of large-scale events on pandemic spread., Comment: 69 pages, 52 figures

Understanding the timing of the peak of a disease outbreak forms an important part of epidemic forecasting. In many cases, such information is essential for planning increased hospital bed demand and for designing of public health interventions. The time taken for an outbreak to become large is inherently stochastic, and therefore uncertain, but after a sufficient number of infections has been reached the subsequent dynamics can be modelled accurately using ordinary differential equations. Here, we present analytical and numerical methods for approximating the time at which a stochastic model of a disease outbreak reaches a large number of cases and for quantifying the uncertainty arising from demographic stochasticity around that time. We then project this uncertainty forwards in time using an ordinary differential equation model in order to obtain a distribution for the peak timing of the epidemic that agrees closely with large simulations but that, for error tolerances relevant to most realistic applications, requires a fraction of the computational cost of full Monte Carlo approaches.

In this paper, we study a two stream species Lotka-Volterra competition patch model with the patches aligned along a line. The two species are supposed to be identical except for the diffusion rates. For each species, the diffusion rates between patches are the same, while the drift rates vary. Our results show that the convexity of the drift rates has a significant impact on the competition outcomes: if the drift rates are convex, then the species with larger diffusion rate wins the competition; if the drift rates are concave, then the species with smaller diffusion rate wins the competition.

In this paper, we study a three-patch two-species Lotka-Volterra competition patch model over a stream network. The individuals are subject to both random and directed movements, and the two species are assumed to be identical except for the movement rates. The environment is heterogeneous, and the carrying capacity is lager in upstream locations. We treat one species as a resident species and investigate whether the other species can invade or not. Our results show that the spatial heterogeneity of environment and the magnitude of the drift rates have a large impact on the competition outcomes of the stream species.

In this paper, we show how to modify a compartmental epidemic model, without changing the dimension, such that separable static heterogeneity is taken into account. The derivation is based on the Kermack–McKendrick renewal equation.

Globally, we are witnessing the rise of complex, non-communicable diseases (NCDs) related to changes in our daily environments. Obesity, asthma, cardiovascular disease, and type 2 diabetes are part of a long list of "lifestyle" diseases that were rare throughout human history but are now common. A key idea from anthropology and evolutionary biology--the evolutionary mismatch hypothesis--seeks to explain this phenomenon. It posits that humans evolved in environments that radically differ from the ones experienced by most people today, and thus traits that were advantageous in past environments may now be "mismatched" and disease-causing. This hypothesis is, at its core, a genetic one: it predicts that loci with a history of selection will exhibit "genotype by environment" (GxE) interactions and have differential health effects in ancestral versus modern environments. Here, we discuss how this concept could be leveraged to uncover the genetic architecture of NCDs in a principled way. Specifically, we advocate for partnering with small-scale, subsistence-level groups that are currently transitioning from environments that are arguably more "matched" with their recent evolutionary history to those that are more "mismatched". These populations provide diverse genetic backgrounds as well as the needed levels and types of environmental variation necessary for mapping GxE interactions in an explicit mismatch framework. Such work would make important contributions to our understanding of environmental and genetic risk factors for NCDs across diverse ancestries and sociocultural contexts.

Previous studies have shown that the topological properties of a complex network, such as heterogeneity and average degree, affect the evolutionary game dynamics on it. However, traditional numerical simulations are usually time-consuming and demand a lot of computational resources. In this paper, we propose the method of dynamical approximate master equations (DAMEs) to accurately approximate the evolutionary outcomes on complex networks. We demonstrate that the accuracy of DAMEs supersedes previous standard pairwise approximation methods, and DAMEs require far fewer computational resources than traditional numerical simulations. We use prisoner's dilemma and snowdrift game on regular and scale-free networks to demonstrate the applicability of DAMEs. Overall, our method facilitates the investigation of evolutionary dynamics on a broad range of complex networks, and provides new insights into the puzzle of cooperation., 21 pages, 4 figures

In varying environments it is beneficial for organisms to utilize available cues to infer the conditions they may encounter and express potentially favorable traits. However, external cues can be unreliable or too costly to use. We consider an alternative strategy where organisms exploit internal sources of information. Even without sensing environmental cues, their internal states may become correlated with the environment as a result of selection, which then form a memory that helps predict future conditions. To demonstrate the adaptive value of such internal memory in varying environments, we revisit the classic example of seed dormancy in annual plants. Previous studies have considered the germination fraction of seeds and its dependence on environmental cues. In contrast, we consider a model of germination fraction that depends on the seed age, which is an internal state that can serve as a memory. We show that, if the environmental variation has temporal structure, then age-dependent germination fractions will allow the population to have an increased long-term growth rate. The more organisms can remember through their internal states, the higher growth rate a population can potentially achieve. Our results suggest experimental ways to infer internal memory and its benefit for adaptation in varying environments., 22 pages, 7 figures

To demonstrate their commitment, for instance during wartime, members of a group will sometimes all engage in a ruinous display. Such widespread, high-cost signals are hard to reconcile with standard models of signaling. For signals to be stable, they must honestly inform their audience, and to be honest, their costs need deter least commited individuals. To account for the existence of uniform high-cost signals , we design a simple game theory model. In our model, senders can engage in second-order signaling. They can pay a cost to express outrage at a non-sender. In doing so, they draw attention to their own signal, and benefit from its increased visibility. Outrage is therefore a self-serving behavior performed at the expense of a target. Using our model and a simulation, we show that outrage can stabilize widespread signals and can lead signal costs to run away. Second-order signaling such as outrage may explain why groups sometimes demand displays of commitment from all their members, and why these displays can entail extreme costs., 11 pages, 4 figures, additional material available at https://evolife.telecom-paris.fr/outrage/RunawaySignals_ArXiv.pdf

Recent studies at individual cell resolution have revealed phenotypic heterogeneity in nominally clonal tumor cell populations. The heterogeneity affects cell growth behaviors, which can result in departure from the idealized exponential growth. Here we measured the stochastic time courses of growth of an ensemble of populations of HL60 leukemia cells in cultures, starting with distinct initial cell numbers to capture the departure from the exponential growth model in the initial growth phase. Despite being derived from the same cell clone, we observed significant variations in the early growth patterns of individual cultures with statistically significant differences in growth kinetics and the presence of subpopulations with different growth rates that endured for many generations. Based on the hypothesis of existence of multiple inter-converting subpopulations, we developed a branching process model that captures the experimental observations.

Maruotti et al. 2022 used a mark-recapture approach to estimate bounds on the true number of monkeypox infections in various countries. We show that such approaches are fundamentally flawed. We argue on conceptual grounds that it is impossible to estimate undercounting based solely on a single stream of reported cases, supporting our contention by applying the authors' method to simulated epidemics. The simulations show that the proposed methods estimate bounds on the ascertainment ratio of ≈0.2-0.5 roughly independently of the true ascertainment ratio. We conclude that the proposed mark-recapture approach should not be used to estimate undercounting or ascertainment ratios. This article is protected by copyright. All rights reserved. Public Health Agency of Canada, NSERC (Discovery grants)

An effective evasion strategy allows prey to survive encounters with predators. Prey are generally thought to escape in a direction that is either random or serves to maximize the minimum distance from the predator. Here we introduce a comprehensive approach to determine the most likely evasion strategy among multiple hypotheses and the role of biomechanical constraints on the escape response of prey fish. Through a consideration of six strategies with sensorimotor noise and previous kinematic measurements, our analysis shows that zebrafish larvae generally escape in a direction orthogonal to the predator's heading. By sensing only the predator's heading, this orthogonal strategy maximizes the distance from fast-moving predators, and, when operating within the biomechanical constraints of the escape response, it provides the best predictions of prey behavior among all alternatives. This work demonstrates a framework for resolving the strategic basis of evastion in predator-prey interactions, which could be applied to a broad diversity of animals., 9 pages, 5 figures

We run stochastic simulations of the spatial version of the rock-paper-scissors game, considering that individuals use sensory abilities to scan the environment to detect the presence of enemies. If the local dangerousness level is above a tolerable threshold, individuals aggregate instead of moving randomly on the lattice. We study the impact of the locally adaptive aggregation on the organisms' spatial organisation by measuring the characteristic length scale of the spatial domains occupied by organisms of a single species. Our results reveal that aggregation is beneficial if triggered when the local density of opponents does not exceed $30\%$; otherwise, the behavioural strategy may harm individuals by increasing the average death risk. We show that if organisms can perceive further distances, they can accurately scan and interpret the signals from the neighbourhood, maximising the effects of the locally adaptive aggregation on the death risk. Finally, we show that the locally adaptive aggregation behaviour promotes biodiversity independently of the organism's mobility. The coexistence probability rises if organisms join conspecifics, even in the presence of a small number of enemies. We verify that our conclusions hold for more complex systems by simulating the generalised rock-paper-scissors models with five and seven species. Our discoveries may be helpful to ecologists in understanding systems where organisms' self-defence behaviour adapts to local environmental cues., 8 pages, 9 figures

Throughout history, prevention and control of dengue transmission have challenged public health authorities worldwide. In the last decades, the interaction of multiple factors, such as environmental and climate variability, has influenced increments in incidence and geographical spread of the virus. In Costa Rica, a country characterized by multiple microclimates separated by short distances, dengue has been endemic since its introduction in 1993. Understanding the role of climatic and environmental factors in the seasonal and inter-annual variability of disease spread is essential to develop effective surveillance and control efforts. In this study, we conducted a wavelet time series analysis of weekly climate, local environmental variables, and dengue cases (2001-2019) from 32 cantons in Costa Rica to identify significant periods (e.g., annual, biannual) in which climate and environmental variables co-varied with dengue cases. Wavelet coherence analysis was used to characterize seasonality, multi-year outbreaks, and relative delays between the time series. Results show that dengue outbreaks occurring every 3 years in cantons located in the country's Central, North, and South Pacific regions were highly coherent with the Oceanic Ni\~no 3.4 and the Tropical North Caribbean Index (TNA). Dengue cases were in phase with El Ni\~no 3.4 and TNA, with El Ni\~no 3.4 ahead of dengue cases by roughly nine months and TNA ahead by less than three months. Annual dengue outbreaks were coherent with local environmental variables (NDWI, EVI, Evapotranspiration, and Precipitation) in most cantons except those located in the Central, South Pacific, and South Caribbean regions of the country. The local environmental variables were in phase with dengue cases and were ahead by around three months., Comment: 21 pages, 15 figures

Mathematical models of genetic evolution often come in pairs, connected by a so-called duality relation. The most seminal example are the Wright-Fisher diffusion and the Kingman coalescent, where the former describes the stochastic evolution of neutral allele frequencies in a large population forward in time, and the latter describes the genetic ancestry of randomly sampled individuals from the population backward in time. As well as providing a richer description than either model in isolation, duality often yields equations satisfied by unknown quantities of interest. We employ the so-called Bernoulli factory, a celebrated tool in simulation-based computing, to derive duality relations for broad classes of genetics models. As concrete examples, we present Wright-Fisher diffusions with general drift functions, and Allen-Cahn equations with general, nonlinear forcing terms. The drift and forcing functions can be interpreted as the action of frequency-dependent selection. To our knowledge, this work is the first time a connection has been drawn between Bernoulli factories and duality in models of population genetics., Comment: 10 pages

With the development of network science, the Turing pattern has been proven to be formed in discrete media such as complex networks, opening up the possibility of exploring it as a generation mechanism in the context of biology, chemistry, and physics. Turing instability in the predator-prey system has been widely studied in recent years. We hope to use the predator-prey interaction relationship in biological populations to explain the influence of network topology on pattern formation. In this paper, we establish a predator-prey model with a weak Allee effect, analyze and verify the Turing instability conditions on the large ER (Erd\"{o}s-R\'{e}nyi) random network with the help of Turing stability theory and numerical experiments, and obtain the Turing instability region. The results show that the diffusion coefficient plays a decisive role in the generation of patterns, and it is interesting that the appropriate initial value will also bring beautiful patterns. When we analyze the model based on the network framework, we find that the average degree of the network has an important impact on the model, and different average degrees will lead to changes in the distribution pattern of the population., Comment: 23 pages, 9 figures, submitted to Nonlinear Analysis: Modelling and Control