arXiv subjects

Simon A. Levin

Publications and source records attributed to Simon A. Levin.

At least 37 records · Page 2Linked to original sources

arXiv2021

sensobol: an R package to compute variance-based sensitivity indices

The R package "sensobol" provides several functions to conduct variance-based uncertainty and sensitivity analysis, from the estimation of sensitivity indices to the visual representation of the results. It implements several state-of-the-art first and total-order estimators and allows the computation of up to third-order effects, as well as of the approximation error, in a swift and user-friendly way. Its flexibility makes it also appropriate for models with either a scalar or a multivariate output. We illustrate its functionality by conducting a variance-based sensitivity analysis of three classic models: the Sobol' (1998) G function, the logistic population growth model of Verhulst (1845), and the spruce budworm and forest model of Ludwig, Jones and Holling (1976).

stat.CO↗

arXiv2021

Phase Transitions and the Theory of Early Warning Indicators for Critical Transitions

Critical transitions, or large changes in the state of a system after a small change in the system's external conditions or parameters, commonly occur in a wide variety of disciplines, from the biological and social sciences to physics. Statistical physics first confronted the problem of emergent phenomena such as critical transitions in the 1800s and 1900s, culminating in the theory of phase transitions. However, although phase transitions show a strong resemblance to critical transitions, the theoretical connections between the two sets of phenomena are tenuous at best, and it would be advantageous to make them more concrete in order to take advantage of the theoretical methods developed by physicists to study phase transitions. Here we attempt to explicitly connect the theory of critical transitions to phase transitions in physics. We initially find something paradoxical, that many critical transitions closely resemble first-order phase transitions, but that many of the early warning indicators developed to anticipate critical transitions, such as critical slowing down or increasing spatial correlations, occur instead in second-order phase transitions. We attempt to reconcile these disparities by making the connection with other phenomena associated with first-order phase transitions, such as spinodal instabilities and metastable states.

cond-mat.stat-mech↗

arXiv2021

A PDE Model for Protocell Evolution and the Origin of Chromosomes via Multilevel Selection

The evolution of complex cellular life involved two major transitions: the encapsulation of self-replicating genetic entities into cellular units and the aggregation of individual genes into a collectively replicating genome. In this paper, we formulate a minimal model of the evolution of proto-chromosomes within protocells. We model a simple protocell composed of two types of genes: a "fast gene" with an advantage for gene-level self-replication and a "slow gene" that replicates more slowly at the gene level, but which confers an advantage for protocell-level reproduction. Protocell-level replication capacity depends on cellular composition of fast and slow genes. We use a partial differential equation to describe how the composition of genes within protocells evolves over time under within-cell and between-cell competition. We find that the gene-level advantage of fast replicators casts a long shadow on the multilevel dynamics of protocell evolution: no level of between-protocell competition can produce coexistence of the fast and slow replicators when the two genes are equally needed for protocell-level reproduction. By introducing a "dimer replicator" consisting of a linked pair of the slow and fast genes, we show analytically that coexistence between the two genes can be promoted in pairwise multilevel competition between fast and dimer replicators, and provide numerical evidence for coexistence in trimorphic competition between fast, slow, and dimer replicators. Our results suggest that dimerization, or the formation of a simple chromosome-like dimer replicator, can help to overcome the shadow of lower-level selection and work in concert with deterministic multilevel selection to allow for the coexistence of two genes that are complementary at the protocell-level but compete at the level of individual gene-level replication.

q-bio.PE↗

arXiv2021

Challenges in cybersecurity: Lessons from biological defense systems

We explore the commonalities between methods for assuring the security of computer systems (cybersecurity) and the mechanisms that have evolved through natural selection to protect vertebrates against pathogens, and how insights derived from studying the evolution of natural defenses can inform the design of more effective cybersecurity systems. More generally, security challenges are crucial for the maintenance of a wide range of complex adaptive systems, including financial systems, and again lessons learned from the study of the evolution of natural defenses can provide guidance for the protection of such systems.

cs.CY↗

arXiv2021

Probabilistic Foundations of Spatial Mean-field Models in Ecology and Applications

Deterministic models of vegetation often summarize, at a macroscopic scale, a multitude of intrinsically random events occurring at a microscopic scale. We bridge the gap between these scales by demonstrating convergence to a mean-field limit for a general class of stochastic models representing each individual ecological event in the limit of large system size. The proof relies on classical stochastic coupling techniques that we generalize to cover spatially extended interactions. The mean-field limit is a spatially extended non-Markovian process characterized by nonlocal integro-differential equations describing the evolution of the probability for a patch of land to be in a given state (the generalized Kolmogorov equations of the process, GKEs). We thus provide an accessible general framework for spatially extending many classical finite-state models from ecology and population dynamics. We demonstrate the practical effectiveness of our approach through a detailed comparison of our limiting spatial model and the finite-size version of a specific savanna-forest model, the so-called Staver-Levin model. There is remarkable dynamic consistency between the GKEs and the finite-size system, in spite of almost sure forest extinction in the finite-size system. To resolve this apparent paradox, we show that the extinction rate drops sharply when nontrivial equilibria emerge in the GKEs, and that the finite-size system's quasi-stationary distribution (stationary distribution conditional on non-extinction) closely matches the bifurcation diagram of the GKEs. Furthermore, the limit process can support periodic oscillations of the probability distribution, thus providing an elementary example of a jump process that does not converge to a stationary distribution. In spatially extended settings, environmental heterogeneity can lead to waves of invasion and front-pinning phenomena.

math.DS↗

arXiv2021

Optimal, near-optimal, and robust epidemic control

In the absence of drugs and vaccines, policymakers use non-pharmaceutical interventions such as social distancing to decrease rates of disease-causing contact, with the aim of reducing or delaying the epidemic peak. These measures carry social and economic costs, so societies may be unable to maintain them for more than a short period of time. Intervention policy design often relies on numerical simulations of epidemic models, but comparing policies and assessing their robustness demands clear principles that apply across strategies. Here we derive the theoretically optimal strategy for using a time-limited intervention to reduce the peak prevalence of a novel disease in the classic Susceptible-Infectious-Recovered epidemic model. We show that broad classes of easier-to-implement strategies can perform nearly as well as the theoretically optimal strategy. But neither the optimal strategy nor any of these near-optimal strategies is robust to implementation error: small errors in timing the intervention produce large increases in peak prevalence. Our results reveal fundamental principles of non-pharmaceutical disease control and expose their potential fragility. For robust control, an intervention must be strong, early, and ideally sustained.

q-bio.PE↗

arXiv2020

Generalized Stoichiometry and Biogeochemistry for Astrobiological Applications

A central need in the field of astrobiology is generalized perspectives on life that make it possible to differentiate abiotic and biotic chemical systems. A key component of many past and future astrobiological measurements is the elemental ratio of various samples. Classic work on Earth's oceans has shown that life displays a striking regularity in the ratio of elements as originally characterized by Redfield. The body of work since the original observations has connected this ratio with basic ecological dynamics and cell physiology, while also documenting the range of elemental ratios found in a variety of environments. Several key questions remain in considering how to best apply this knowledge to astrobiological contexts: How can the observed variation of the elemental ratios be more formally systematized using basic biological physiology and ecological or environmental dynamics? How can these elemental ratios be generalized beyond the life that we have observed on our own planet? Here we expand recently developed generalized physiological models to create a simple framework for predicting the variation of elemental ratios found in various environments. We then discuss further generalizing the physiology for astrobiological applications. Much of our theoretical treatment is designed for in situ measurements applicable to future planetary missions. We imagine scenarios where three measurements can be made - particle/cell sizes, particle/cell stoichiometry, and fluid or environmental stoichiometry - and develop our theory in connection with these often deployed measurements.

q-bio.QM↗

arXiv2020

A well-timed switch from local to global agreements accelerates climate change mitigation

Recent attempts at cooperating on climate change mitigation highlight the limited efficacy of large-scale agreements, when commitment to mitigation is costly and initially rare. Bottom-up approaches using region-specific mitigation agreements promise greater success, at the cost of slowing global adoption. Here, we show that a well-timed switch from regional to global negotiations dramatically accelerates climate mitigation compared to using only local, only global, or both agreement types simultaneously. This highlights the scale-specific roles of mitigation incentives: local incentives capitalize on regional differences (e.g., where recent disasters incentivize mitigation) by committing early-adopting regions, after which global agreements draw in late-adopting regions. We conclude that global agreements are key to overcoming the expenses of mitigation and economic rivalry among regions but should be attempted once regional agreements are common. Gradually up-scaling efforts could likewise accelerate mitigation at smaller scales, for instance when costly ecosystem restoration initially faces limited public and legislative support.

nlin.AO↗

arXiv2020

Active Control and Sustained Oscillations in actSIS Epidemic Dynamics

An actively controlled Susceptible-Infected-Susceptible (actSIS) contagion model is presented for studying epidemic dynamics with continuous-time feedback control of infection rates. Our work is inspired by the observation that epidemics can be controlled through decentralized disease-control strategies such as quarantining, sheltering in place, social distancing, etc., where individuals actively modify their contact rates with others in response to observations of infection levels in the population. Accounting for a time lag in observations and categorizing individuals into distinct sub-populations based on their risk profiles, we show that the actSIS model manifests qualitatively different features as compared with the SIS model. In a homogeneous population of risk-averters, the endemic equilibrium is always reduced, although the transient infection level can exhibit overshoot or undershoot. In a homogeneous population of risk-tolerating individuals, the system exhibits bistability, which can also lead to reduced infection. For a heterogeneous population comprised of risk-tolerators and risk-averters, we prove conditions on model parameters for the existence of a Hopf bifurcation and sustained oscillations in the infected population.

physics.soc-ph↗

arXiv2020

Staggered Release Policies for COVID-19 Control: Costs and Benefits of Sequentially Relaxing Restrictions by Age

Strong social distancing restrictions have been crucial to controlling the COVID-19 outbreak thus far, and the next question is when and how to relax these restrictions. A sequential timing of relaxing restrictions across groups is explored in order to identify policies that simultaneously reduce health risks and economic stagnation relative to current policies. The goal will be to mitigate health risks, particularly among the most fragile sub-populations, while also managing the deleterious effect of restrictions on economic activity. The results of this paper show that a properly constructed sequential release of age-defined subgroups from strict social distancing protocols can lead to lower overall fatality rates than the simultaneous release of all individuals after a lockdown. The optimal release policy, in terms of minimizing overall death rate, must be sequential in nature, and it is important to properly time each step of the staggered release. This model allows for testing of various timing choices for staggered release policies, which can provide insights that may be helpful in the design, testing, and planning of disease management policies for the ongoing COVID-19 pandemic and future outbreaks.

q-bio.PE↗

arXiv2019

Coalition-structured governance improves cooperation to provide public goods

While the benefits of common and public goods are shared, they tend to be scarce when contributions are provided voluntarily. Failure to cooperate in the provision or preservation of these goods is fundamental to sustainability challenges, ranging from local fisheries to global climate change. In the real world, such cooperative dilemmas occur in multiple interactions with complex strategic interests and frequently without full information. We argue that voluntary cooperation enabled across multiple coalitions (akin to polycentricity) not only facilitates greater generation of non-excludable public goods, but may also allow evolution toward a more cooperative, stable, and inclusive approach to governance. Contrary to any previous study, we show that these merits of multi-coalition governance are far more general than the singular examples occurring in the literature, and are robust under diverse conditions of excludability, congestability of the non-excludable public good, and arbitrary shapes of the return-to-contribution function. We first confirm the intuition that a single coalition without enforcement and with players pursuing their self-interest without knowledge of returns to contribution is prone to cooperative failure. Next, we demonstrate that the same pessimistic model but with a multi-coalition structure of governance experiences relatively higher cooperation by enabling recognition of marginal gains of cooperation in the game at stake. In the absence of enforcement, public-goods regimes that evolve through a proliferation of voluntary cooperative forums can maintain and increase cooperation more successfully than singular, inclusive regimes.

econ.GN↗

arXiv2019

Consensus and Polarisation in Competing Complex Contagion Processes

The rate of adoption of new information depends on reinforcement from multiple sources in a way that often cannot be described by simple contagion processes. In such cases, contagion is said to be complex. Complex contagion happens in the diffusion of human behaviours, innovations, and knowledge. Based on that evidence, we propose a model that considers multiple, potentially asymmetric, and competing contagion processes and analyse its respective population-wide dynamics, bringing together ideas from complex contagion, opinion dynamics, evolutionary game theory, and language competition by shifting the focus from individuals to the properties of the diffusing processes. We show that our model spans a dynamical space in which the population exhibits patterns of consensus, dominance, and, importantly, different types of polarisation, a more diverse dynamical environment that contrasts with single simple contagion processes. We show how these patterns emerge and how different population structures modify them through a natural development of spatial correlations: structured interactions increase the range of the dominance regime by reducing that of dynamic polarisation, tight modular structures can generate structural polarisation, depending on the interplay between fundamental properties of the processes and the modularity of the interaction network.

physics.soc-ph↗

arXiv2019

Bacteria push the limits of chemotactic precision to navigate dynamic chemical gradients

Ephemeral aggregations of bacteria are ubiquitous in the environment, where they serve as hotbeds of metabolic activity, nutrient cycling, and horizontal gene transfer. In many cases, these regions of high bacterial concentration are thought to form when motile cells use chemotaxis to navigate to chemical hotspots. However, what governs the dynamics of bacterial aggregations is unclear. Here, we use a novel experimental platform to create realistic sub-millimeter scale nutrient pulses with controlled nutrient concentrations. By combining experiments, mathematical theory and agent-based simulations, we show that individual \textit{Vibrio ordalii} bacteria begin chemotaxis toward hotspots of dissolved organic matter (DOM) when the magnitude of the chemical gradient rises sufficiently far above the sensory noise that is generated by stochastic encounters with chemoattractant molecules. Each DOM hotspot is surrounded by a dynamic ring of chemotaxing cells, which congregate in regions of high DOM concentration before dispersing as DOM diffuses and gradients become too noisy for cells to respond to. We demonstrate that \textit{V. ordalii} operates close to the theoretical limits on chemotactic precision. Numerical simulations of chemotactic bacteria, in which molecule counting noise is explicitly taken into account, point at a tradeoff between nutrient acquisition and the cost of chemotactic precision. More generally, our results illustrate how limits on sensory precision can be used to understand the location, spatial extent, and lifespan of bacterial behavioral responses in ecologically relevant environments.

physics.bio-ph↗

arXiv2015

Physical Limits on Bacterial Navigation in Dynamic Environments

Many chemotactic bacteria inhabit environments in which chemicals appear as localized pulses and evolve by processes such as diffusion and mixing. We show that, in such environments, physical limits on the accuracy of temporal gradient sensing govern when and where bacteria can accurately measure the cues they use to navigate. Chemical pulses are surrounded by a predictable dynamic region, outside which bacterial cells cannot resolve gradients above noise. The outer boundary of this region initially expands in proportion to $\sqrt{t}$, before rapidly contracting. Our analysis also reveals how chemokinesis - the increase in swimming speed many bacteria exhibit when absolute chemical concentration exceeds a threshold - may serve to enhance chemotactic accuracy and sensitivity when the chemical landscape is dynamic. More generally, our framework provides a rigorous method for partitioning bacteria into populations that are "near" and "far" from chemical hotspots in complex, rapidly evolving environments such as those that dominate aquatic ecosystems.

physics.bio-ph↗

arXiv2013

Evolutionary comparison between viral lysis rate and latent period

Marine viruses shape the structure of the microbial community. They are, thus, a key determinant of the most important biogeochemical cycles in the planet. Therefore, a correct description of the ecological and evolutionary behavior of these viruses is essential to make reliable predictions about their role in marine ecosystems. The infection cycle, for example, is indistinctly modeled in two very different ways. In one representation, the process is described including explicitly a fixed delay between infection and offspring release. In the other, the offspring are released at exponentially distributed times according to a fixed release rate. By considering obvious quantitative differences pointed out in the past, the latter description is widely used as a simplification of the former. However, it is still unclear how the dichotomy "delay versus rate description" affects long-term predictions of host-virus interaction models. Here, we study the ecological and evolutionary implications of using one or the other approaches, applied to marine microbes. To this end, we use mathematical and eco-evolutionary computational analysis. We show that the rate model exhibits improved competitive abilities from both ecological and evolutionary perspectives in steady environments. However, rate-based descriptions can fail to describe properly long-term microbe-virus interactions. Moreover, additional information about trade-offs between life-history traits is needed in order to choose the most reliable representation for oceanic bacteriophage dynamics. This result affects deeply most of the marine ecosystem models that include viruses, especially when used to answer evolutionary questions.

q-bio.PE↗

arXiv2012

Patchiness and Demographic Noise in Three Ecological Examples

Understanding the causes and effects of spatial aggregation is one of the most fundamental problems in ecology. Aggregation is an emergent phenomenon arising from the interactions between the individuals of the population, able to sense only -at most- local densities of their cohorts. Thus, taking into account the individual-level interactions and fluctuations is essential to reach a correct description of the population. Classic deterministic equations are suitable to describe some aspects of the population, but leave out features related to the stochasticity inherent to the discreteness of the individuals. Stochastic equations for the population do account for these fluctuation-generated effects by means of demographic noise terms but, owing to their complexity, they can be difficult (or, at times, impossible) to deal with. Even when they can be written in a simple form, they are still difficult to numerically integrate due to the presence of the "square-root" intrinsic noise. In this paper, we discuss a simple way to add the effect of demographic stochasticity to three classic, deterministic ecological examples where aggregation plays an important role. We study the resulting equations using a recently-introduced integration scheme especially devised to integrate numerically stochastic equations with demographic noise. Aimed at scrutinizing the ability of these stochastic examples to show aggregation, we find that the three systems not only show patchy configurations, but also undergo a phase transition belonging to the directed percolation universality class.

q-bio.PE↗

arXiv2012

Multiscale analysis of collective motion and decision-making in swarms: An advection-diffusion equation with memory approach

We propose a (time) multiscale method for the coarse-grained analysis of self--propelled particle models of swarms comprising a mixture of `naïve' and `informed' individuals, used to address questions related to collective motion and collective decision--making in animal groups. The method is based on projecting the particle configuration onto a single `meta-particle' that consists of the group elongation and the mean group velocity and position. The collective states of the configuration can be associated with the transient and asymptotic transport properties of the random walk followed by the meta-particle. These properties can be accurately predicted by an advection-diffusion equation with memory (ADEM) whose parameters are obtained from a mean group velocity time series obtained from a single simulation run of the individual-based model.

q-bio.PE↗

arXiv2011

Evolution of a Modular Software Network

"Evolution behaves like a tinkerer" (Francois Jacob, Science, 1977). Software systems provide a unique opportunity to understand biological processes using concepts from network theory. The Debian GNU/Linux operating system allows us to explore the evolution of a complex network in a novel way. The modular design detected during its growth is based on the reuse of existing code in order to minimize costs during programming. The increase of modularity experienced by the system over time has not counterbalanced the increase in incompatibilities between software packages within modules. This negative effect is far from being a failure of design. A random process of package installation shows that the higher the modularity the larger the fraction of packages working properly in a local computer. The decrease in the relative number of conflicts between packages from different modules avoids a failure in the functionality of one package spreading throughout the entire system. Some potential analogies with the evolutionary and ecological processes determining the structure of ecological networks of interacting species are discussed.

cs.OS↗