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Populations and Evolution

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preprint2026arXiv

Conditioning as a route to stereotyped behavior in growing populations

Biological systems perform complex multi-step processes in a reproducible way despite underlying stochasticity. The standard explanation is micromanagement by molecular machinery that recognizes and corrects specific errors. Here we study conditioning, a qualitatively different strategy in which attempts failing a coarse criterion are destroyed and do not leave a physical record. The surviving, i.e., conditioned, ensemble is narrower and therefore more ordered. We model conditioning through stochastic resets in a ''socks-before-shoes'' model of a growing population, where $n$ actions must be completed in any order to replicate and any replication attempt not finished by a threshold time is discarded. We find that resets impose hierarchical temporal ordering of the $n$ actions without microscopic control over which action happens when. When disorder carries a sufficient time penalty, this ordering is free: the fastest-growing population is automatically the most ordered, with no direct selection for order required. Save points, at which verified progress is preserved across resets, allow conditioning to scale to complex multi-step processes. Conditioning provides a minimal route to reliable behavior, requiring only a clock rather than molecular machinery that recognizes specific errors. For the right class of processes, it pays for itself.

preprint2022arXiv

Evaluation of non-pharmaceutical interventions and optimal strategies for containing the COVID-19 pandemic

Given multiple new COVID-19 variants are continuously emerging, non-pharmaceutical interventions are still primary control strategies to curb the further spread of coronavirus. However, implementing strict interventions over extended periods of time is inevitably hurting the economy. With an aim to solve this multi-objective decision-making problem, we investigate the underlying associations between policies, mobility patterns, and virus transmission. We further evaluate the relative performance of existing COVID-19 control measures and explore potential optimal strategies that can strike the right balance between public health and socio-economic recovery for individual states in the US. The results highlight the power of state of emergency declaration and wearing face masks and emphasize the necessity of pursuing tailor-made strategies for different states and phases of epidemiological transmission. Our framework enables policymakers to create more refined designs of COVID-19 strategies and can be extended to inform policy makers of any country about best practices in pandemic response.

preprint2022arXiv

Efficient Stochastic Simulation of Network Topology Effects on the Peak Number of Infections in Epidemic Outbreaks

This paper investigates the effect of the structure of the contact network on the dynamics of the epidemic outbreak. In particular, we focus on the peak number of critically infected nodes (PCIN), determining the maximum workload of intensive healthcare units which should be kept low. As a model and simulation method, we develop a continuous-time Markov chain (CTMC) model and an efficient simulation-based on Gillespie's Stochastic Simulation Algorithm (SSA). This methods combine a realistic approximation of the stochastic process not relying on the assumptions of mean-field models and large asymptotically large population sizes as in differential equation models, and at the same time an efficient way to simulate networks of moderate size. The approach is analysed for different scenarios, based on data from the COVID-19 outbreak and demographic data from Ukraine. From these results we extract network topology features that need to be considered to effectively decrease the peak number of infections. The CTMC simulation is implemented in python and integrated in a dashboard that can be used for interactive exploration and it is made openly available.

preprint2026arXiv

Phylogenetic Tree Inference with Tropical Axial Attention

In this work, we introduce a Tropical Axial Attention neural reasoning architecture that replaces vanilla softmax dot-product attention with max-plus operators, inducing a piecewise-linear structure aligned with dynamic programming formulations. From multi-species sequence alignments, our model learns all possible pairwise distances and is trained using a combination of $\ell_1$ and tropical symmetric distance metric losses with an ultrametric violation penalty. We leverage the well known isomorphic relationship between the space of all phylogenetic trees with $n$ species and tropical Grassmannian to show that tropical attention provides a natural geometric framework for phylogenetic inference. On empirical $DS1-DS11$ alignments, where true trees are unknown, the tropical model produces distance matrices that are substantially closer to their BME-induced tree metrics than the baseline models. These results suggest that tropical attention is a useful geometric inductive bias for neural phylogenetic inference, especially under distribution shift and when tree-metric consistency is important.

preprint2022arXiv

Predator-Prey Linear Coupling with Hybrid Species

The classical two-species non-linear Predator-Prey system, often used in population dynamics modeling, is expressed in terms of a single positive coupling parameter $λ$. Based on standard logarithmic transformations, we derive a novel $λ$-\textit{invariant} Hamiltonian resulting in two coupled first-order ODEs for ``hybrid-species'', \textit{albeit} with one being \textit{linear}; we thus derive a new exact, closed-form, single quadrature solution valid for any value of $λ$ and the system's energy. In the particular case $λ= 1$ the ODE system completely uncouples and a new, exact, energy-only dependent simple quadrature solution is derived. In the case $λ\neq 1$ an accurate practical approximation uncoupling the non-linear system is proposed and solutions are provided in terms of explicit quadratures together with high energy asymptotic solutions. A novel, exact, closed-form expression of the system's oscillation period valid for any value of $λ$ and orbital energy is also derived; two fundamental properties of the period are established; for $λ= 1$ the period is expressed in terms of a universal energy function and shown to be the shortest.

preprint2026arXiv

Travelling waves of invasion in microbial communities with phenotypic switching

Complex microbial habitats see the spatial competition of different clonal bacterial populations that switch between different phenotypes. Here, we determine the effect of this subpopulation structure on the invasion of one species by another in a minimal model of two competing species: one species switches, both stochastically and in response to its competitor, to a persister phenotype resilient to competition. Surprisingly, our combined analytical and numerical results show that this phenotypic switching has no effect on the speed of the travelling wave by which the competitors invade the first population. Conversely, we discover that phenotypic switching can speed up the wave by which this population invades their competitors. Our results thus suggest, counterintuitively, that bacterial persistence can be an offensive, rather than defensive ecological strategy.

preprint2022arXiv

Studying the mixed transmission in a community with age heterogeneity: COVID-19 as a case study

COVID-19 has been prevalent worldwide for about 2 years now and has brought unprecedented challenges to our society. Before vaccines were available, the main disease intervention strategies were non-pharmaceutical. Starting December 2020, in Ontario, Canada, vaccines were approved for administering to vulnerable individuals and gradually expanded to all individuals above the age of 12. As the vaccine coverage reached a satisfactory level among the eligible population, normal social activities resumed and schools reopened starting September 2021. However, when schools reopen for in-person learning, children under the age of 12 are unvaccinated and are at higher risks of contracting the virus. We propose an age-stratified model based on the age and vaccine eligibility of the individuals. We fit our model to the data in Ontario, Canada and obtain a good fitting result. The results show that a relaxed between-group contact rate may trigger future epidemic waves more easily than an increased within-group contact rate. An increasing mixed contact rate of the older group quickly amplifies the daily incidence numbers for both groups whereas an increasing mixed contact rate of the younger group mainly leads to future waves in the younger group alone. The results indicate the importance of accelerating vaccine rollout for younger individuals in mitigating disease spread.

preprint2026arXiv

Direct From Darwin: Deriving Advanced Optimizers From Evolutionary First Principles

Evolutionary computation has long promised to deliver both high-performance optimization tools as well as rigorous scientific simulations of Darwinian evolution. However, modern algorithms frequently abandon evolutionary fidelity for physics-inspired heuristics or superficial biological metaphors. This paper derives a suite of advanced gradient-based optimization algorithms directly from evolutionary first principles. We introduce Darwinian Lineage Simulations (DLS) to prove that, in an asexual context, Fisher's and Wright's historically opposed views of evolution are actually formally equivalent; One can partition Fisher's deterministically-evolving total population into Wright's randomly-drifting sub-populations. We prove that proper bookkeeping requires introducing a specific kind of structured noise (the DLS noise relation). Crucially, any bookkeeping choices which satisfy this relation will yield a faithful simulation of evolution. Using this vast representational freedom, we prove that a broad family of battle-tested optimization algorithms are already perfectly compatible with evolutionary dynamics. These include: Stochastic Gradient Descent as well as many regularizations/approximations of Newton's method and Natural Gradient Descent. By simply adding DLS noise (i.e., evolutionarily faithful genetic drift), these algorithms become scientifically valid in silico simulations of Darwinian evolution. Finally, we demonstrate that even the state-of-the-art Adam optimizer can be brought into evolutionary compliance through a minor mathematical surgery.

preprint2022arXiv

Designing weights for quartet-based methods when data is heterogeneous across lineages

Homogeneity across lineages is a common assumption in phylogenetics according to which nucleotide substitution rates remain constant in time and do not depend on lineages. This is a simplifying hypothesis which is often adopted to make the process of sequence evolution more tractable. However, its validity has been explored and put into question in several papers. On the other hand, dealing successfully with the general case (heterogeneity across lineages) is one of the key features of phylogenetic reconstruction methods based on algebraic tools. The goal of this paper is twofold. First, we present a new weighting system for quartets (ASAQ) based on algebraic and semi-algebraic tools, thus specially indicated to deal with data evolving under heterogeneus rates. This method combines the weights two previous methods by means of a test based on the positivity of the branch length estimated with the paralinear distance. ASAQ is statistically consistent when applied to GM data, considers rate and base composition heterogeneity among lineages and does not assume stationarity nor time-reversibility. Second, we test and compare the performance of several quartet-based methods for phylogenetic tree reconstruction (namely, Quartet Puzzling, Weight Optimization and Wilson's method) in combination with ASAQ weights and other weights based on algebraic and semi-algebraic methods or on the paralinear distance. These tests are applied to both simulated and real data and support Weight Optimization with ASAQ weights as a reliable and successful reconstruction method.

preprint2025arXiv

Quenched coalescent for diploid population models with selfing and overlapping generations

We introduce a general diploid population model with self-fertilization and possible overlapping generations, and study the genealogy of a sample of $n$ genes as the population size $N$ tends to infinity. Unlike traditional approach in coalescent theory which considers the unconditional (annealed) law of the gene genealogies averaged over the population pedigree, here we study the conditional (quenched) law of gene genealogies given the pedigree. We focus on the case of high selfing probability and obtain that this conditional law converges to a random probability measure, given by the random law of a system of coalescing random walks on an exchangeable fragmentation-coalescence process of \cite{berestycki04}. This system contains the system of coalescing random walks on the ancestral recombination graph as a special case, and it sheds new light on the site-frequency spectrum (SFS) of genetic data by specifying how SFS depends on the pedigree. The convergence result is proved by means of a general characterization of weak convergence for random measures on the Skorokhod space with paths taking values in a locally compact Polish space.

preprint2025arXiv

Binary Galton-Watson trees with mutations

We consider a multitype Galton-Watson process that allows for the mutation and reversion of individual types in discrete and continuous time. In this setting, we explicitly compute the time evolution of quantities such as the mean and distributions of different types. This allows us in particular to estimate the proportions of different types in the long run, as well as the distribution of the first time of occurrence of a given type as the tree size or time increases. Our approach relies on the recursive computation of the joint distribution of types conditionally to the value of the total progeny. In comparison with the literature on related multitype models, we do not rely on approximations.

preprint2022arXiv

Transport and Optimal Control of Vaccination Dynamics for COVID-19

We develop a mathematical model for transferring the vaccine BNT162b2 based on the heat diffusion equation. Then, we apply optimal control theory to the proposed generalized SEIR model. We introduce vaccination for the susceptible population to control the spread of the COVID-19 epidemic. For this, we use the Pontryagin minimum principle to find the necessary optimality conditions for the optimal control. The optimal control problem and the heat diffusion equation are solved numerically. Finally, several simulations are done to study and predict the spread of the COVID-19 epidemic in Italy. In particular, we compare the model in the presence and absence of vaccination.

preprint2022arXiv

Estimates of solutions in a model of antiviral immune response

We consider a model of antiviral immune response proposed in the works of G.I. Marchuk. The model is described by a system of delay differential equations. The asymptotic stability of a stationary solution to the system corresponding to a completely healthy organism is studied. Estimates of the attraction set of the given stationary solution are obtained and estimates of solutions characterizing the stabilization rate at infinity are established. The results are obtained using the Lyapunov-Krasovskii functional.

preprint2022arXiv

Computational Modelling of Plasticity-Led Evolution

Plasticity-led evolution is a form of evolution where a change in the environment induces novel traits via phenotypic plasticity, after which the novel traits are genetically accommodated over generations under the novel environment. This mode of evolution is expected to resolve the problem of gradualism (i.e., evolution by the slow accumulation of mutations that induce phenotypic variation) implied by the Modern Evolutionary Synthesis, in the face of a large environmental change. While experimental works are essential for validating that plasticity-led evolution indeed happened, we need computational models to gain insight into its underlying mechanisms and make qualitative predictions. Such computational models should include the developmental process and gene-environment interactions in addition to genetics and natural selection. We point out that gene regulatory network models can incorporate all the above notions. In this review, we highlight results from computational modelling of gene regulatory networks that consolidate the criteria of plasticity-led evolution. Since gene regulatory networks are mathematically equivalent to artificial recurrent neural networks, we also discuss their analogies and discrepancies, which may help further understand the mechanisms underlying plasticity-led evolution.

preprint2026arXiv

Essential Role of Extrinsic Noise in Models of E. coli Division Control

Our understanding of cell division control in bacteria still relies largely on interpreting correlations between phenomenological variables, with limited connection to the underlying molecular mechanisms. Here, we analytically solve a stochastic threshold-accumulation model in which a size-dependent divisor protein triggers division upon reaching a noisy, autocorrelated threshold, quantifying within a unified framework the combined effects of intrinsic and extrinsic noise and key mechanistic parameters such as protein reset and threshold memory. We show that incorporating these elements yields behavior far richer than the commonly assumed adder, spanning a continuum of division strategies from timer to sizer while modulating size fluctuations in a nontrivial fashion. Comparison with single-cell E. coli data shows that extrinsic noise and additional mechanistic ingredients are required to account for the observed size fluctuations. The adder emerges when threshold correlations balance protein reset, generalizing the hypothesis that full reset is necessary to maintain adder control. Our results establish a unified analytical framework linking stochastic molecular processes to emergent division laws, to be used in more complex bacterial cell-cycle models.

preprint2022arXiv

Host movement, transmission hot spots, and vector-borne disease dynamics on spatial networks

We examine how spatial heterogeneity combines with mobility network structure to influence vector-borne disease dynamics. Specifically, we consider a Ross-Macdonald-type disease model on $n$ spatial locations that are coupled by host movement on a strongly connected, weighted, directed graph. We derive a closed form approximation to the domain reproduction number using a Laurent series expansion, and use this approximation to compute sensitivities of the basic reproduction number to model parameters. To illustrate how these results can be used to help inform mitigation strategies, as a case study we apply these results to malaria dynamics in Namibia, using published cell phone data and estimates for local disease transmission. Our analytical results are particularly useful for understanding drivers of transmission when mobility sinks and transmission hot spots do not coincide.

preprint2015arXiv

Individual-based approach to epidemic processes on arbitrary dynamic contact networks

The dynamics of contact networks and epidemics of infectious diseases often occur on comparable time scales. Ignoring one of these time scales may provide an incomplete understanding of the population dynamics of the infection process. We develop an individual-based approximation for the susceptible-infected-recovered epidemic model applicable to arbitrary dynamic networks. Our framework provides, at the individual-level, the probability flow over time associated with the infection dynamics. This computationally efficient framework discards the correlation between the states of different nodes, yet provides accurate results in approximating direct numerical simulations. It naturally captures the temporal heterogeneities and correlations of contact sequences, fundamental ingredients regulating the timing and size of an epidemic outbreak. Using real-life data, we show that the static network model overestimates the reproduction number but underestimates the infection potential of super-spreading individuals. The high accuracy of our approximation further allows us to detect the index individual of an epidemic outbreak.

preprint2024arXiv

Dynamic coexistence driven by physiological transitions in microbial communities

Microbial ecosystems are commonly modeled by fixed interactions between species in steady exponential growth states. However, microbes often modify their environments so strongly that they are forced out of the exponential state into stressed or non-growing states. Such dynamics are typical of ecological succession in nature and serial-dilution cycles in the laboratory. Here, we introduce a phenomenological model, the Community State model, to gain insight into the dynamic coexistence of microbes due to changes in their physiological states. Our model bypasses specific interactions (e.g., nutrient starvation, stress, aggregation) that lead to different combinations of physiological states, referred to collectively as "community states", and modeled by specifying the growth preference of each species along a global ecological coordinate, taken here to be the total community biomass density. We identify three key features of such dynamical communities that contrast starkly with steady-state communities: increased tolerance of community diversity to fast growth rates of species dominating different community states, enhanced community stability through staggered dominance of different species in different community states, and increased requirement on growth dominance for the inclusion of late-growing species. These features, derived explicitly for simplified models, are proposed here to be principles aiding the understanding of complex dynamical communities. Our model shifts the focus of ecosystem dynamics from bottom-up studies based on idealized inter-species interaction to top-down studies based on accessible macroscopic observables such as growth rates and total biomass density, enabling quantitative examination of community-wide characteristics.

preprint2026arXiv

Genotype specificity and spatial arrangement govern the direction and magnitude of selection in variable environments

Spatial environmental variation can either amplify or suppress the fixation of beneficial mutants in structured populations, yet the interplay of ecological factors and spatial structure in determining which outcome occurs remains theoretically unresolved. Here, we develop a unified framework for selection on lattice graphs with environmental heterogeneity, in which mutant and resident fitness depend on the local environmental state. Across three common classes of genotype-environment interactions and a wide range of spatial arrangements of environmental states, we identify two governing principles. Genotype specificity determines the direction of the effect: heterogeneity amplifies selection when it modulates resident fitness, but suppresses selection when it modulates mutant fitness, with genotype-symmetric modulation producing weaker amplification. Spatial arrangement determines the magnitude: intermixed versus clustered environments tune the strength of amplification or suppression without reversing the direction of the effect. Together, these principles reconcile disparate theoretical results and provide predictive criteria for adaptation in heterogeneous landscapes, from microbial communities to somatic evolution and cancer.

preprint2026arXiv

Tree-Conditioned Edit Flows for Ancestral Sequence Reconstruction

Ancestral sequence reconstruction (ASR) aims to infer extinct protein sequences at internal nodes of a phylogenetic tree. Classical ASR methods are typically based on continuous-time Markov substitution models, but they treat sites largely independently and handle insertions and deletions only weakly or not at all. We introduce a tree-conditioned edit-flow model for variable-length ASR. Given two descendant sequences and their branch distances to a shared ancestor, the model reconstructs the ancestor through paired bidirectional edit trajectories constrained to agree on a common ancestral state. On a benchmark of experimentally evolved sequences with only context-independent substitutions, the model does not match the accuracy of the best classical method, yet still achieves reasonable performance despite being trained on natural sequences that include insertions, deletions, and substitutions. On a benchmark of natural homologous sequences with abundant insertions and deletions, the model most accurately localizes inferred evolutionary change.

preprint2022arXiv

The effect of biologically mediated decay rates on modelling soil carbon sequestration in agricultural settings

Microbial biomass carbon (MBC), a crucial soil labile carbon fraction, is the most active component of the soil organic carbon (SOC) that regulates bio-geochemical processes in terrestrial ecosystems. Some studies in the literature ignore the effect of microbial population growth on carbon decomposition rates. In reality, we might expect that the decomposition rate should be related to the population of microbes in the soil and have a positive relationship with the size of the microbial biomass pool. In this study, we explore the effect of microbial population growth on the accuracy of modelling soil carbon sequestration by developing and comparing two soil carbon models that consider a carrying capacity and limit to the growth of the microbial pool. We apply our models to three datasets, two small and one large datasets, and we select the best model in terms of having the best predictive performance through two model selection methods. Through this analysis we reveal that commonly used complex soil carbon models can over-fit in the presence of both small and large time-series datasets, and our simpler model can produce more accurate predictions. We conclude that considering the microbial population growth in a soil carbon model improves the accuracy of a model in the presence of a large dataset.

preprint2026arXiv

EPITIME: A Computational Framework for Integral Epidemic Models with Structure-Preserving Discretizations

We present EPITIME (EPidemic Integral models TIMe profile Explorer), a computational framework for the simulation of two classes of integral epidemic models: an age of infection model and an information dependent behavioural model. The framework combines structure preserving Non-Standard Finite Difference discretizations with modular implementations in MATLAB and Python, together with routines for parameter handling, input validation, performance assessment, and graphical interaction. The proposed methods preserve key qualitative properties of the continuous problems, including positivity, boundedness, invariant regions, and correct long term behaviour, independently of the time step. We outline the numerical schemes for both model classes and their main analytical properties, including first order convergence. We then describe the software architecture and illustrate its use through numerical experiments on asymptotic behaviour, inverse reconstruction of an infectivity kernel from COVID 19 incidence data, and behavioural dynamics under different memory kernels. Overall, EPITIME provides a reliable and accessible computational environment for the numerical study of renewal epidemic models.

preprint2022arXiv

From Microscopic SDE to Stochastic Macroscopic Equation: A Framework for Modeling Diversity in Cancer and Other Complex Living Systems

Biological living systems in general exhibit complex and diverse dynamics. The latter, in particular, is essential, since diversification increases the odds of survival of an organism while reducing the risk of extinction of the population. Primarily, diversification is a consequence of the randomness in the replication process of a biological cell, which eventually manifests into a motley set of macroscopic features of an individual. These heterogeneous features of individuals constitutes for diversity in population. Cancer is a prime example of such a complex system where the transformed cells exhibit plethora of disparate features, which in turn makes modeling their dynamics quite challenging. In this paper we consider cancer as a prototype of a complex living system and provide two contrasting perspective for studying and modeling it. Based on this we illicit a deeper role of diversification in the evolution of cancer. Following this, we ask ourselves how can we model these diverse dynamics in a multiscale setting. We address the shortcoming of the existing multiscale modeling technique by providing an abstract but mathematically rigorous framework for deducing stochastic evolution equations at the macroscopic level starting from a microscopic description of the involved dynamics. We achieve this by making use of the connection between stochastic process and the semigroup operator generated by them. In particular, we look at the semigroups generated by Levy processes and their connection with the characteristic functions and Levy symbols. The latter turns out to represent pseudo-differential operators using which we eventually provide a mechanism for constructing stochastic evolution equations. Altogether, this provides the framework for modeling diverse dynamics at the macroscale starting from the microscale.

preprint2022arXiv

Structured interactions as a stabilizing mechanism for competitive ecosystems

How large ecosystems can create and maintain the remarkable biodiversity we see in nature is probably one of the biggest open questions in science, attracting attention from different fields, from Theoretical Ecology to Mathematics and Physics. In this context, modeling the stable coexistence of species competing for limited resources is a particularly challenging task. From a mathematical point of view, coexistence in competitive dynamics can be achieved when dominance among species forms intransitive loops. However, these relationships usually lead to species' relative abundances neutrally cycling without converging to a stable equilibrium. Although in recent years several mechanisms have been proposed, models able to explain species coexistence in competitive communities are still limited. Here we identify locality in the interactions as one of the simplest mechanisms leading to stable species coexistence. We consider a simplified ecosystem where individuals of each species lay on a spatial network and interactions are possible only between nodes within a certain distance. Varying such distance allows to interpolate between local and global competition. Our results demonstrate, within the scope of our model, that species coexist reaching a stable equilibrium when two conditions are met: individuals are embedded in space and can only interact with other individuals within a short distance. On the contrary, when one of these ingredients is missing, large oscillations and neutral cycles emerge.