Stochastic effects on the dynamics of an epidemic due to population subdivision
Philip Bittihn, Ramin Golestanian
备注：
A proposal for a containment/exit strategy in response to the COVID-19 pandemic. Note the change in title from the original (2020/3/19) version "Containment strategy for an epidemic based on fluctuations in the SIR model"
Using a stochastic Susceptible-Infected-Removed (SIR) meta-population model of disease transmission, we present analytical calculations and numerical simulations dissecting the interplay between stochasticity and the division of a population into mutually independent sub-populations. We show that subdivision activates two stochastic effects---extinction and desynchronization---diminishing the overall impact of the outbreak, even when the total population has already left the stochastic regime and the basic reproduction number is not altered by the subdivision. Both effects are quantitatively captured by our theoretical estimates, allowing us to determine their individual contributions to the observed reduction of the peak of the epidemic.
Natural selection drives populations towards higher fitness, but crossing fitness valleys or plateaus may facilitate progress up a rugged fitness landscape involving epistasis. We investigate quantitatively the effect of subdividing an asexual population on the time it takes to cross a fitness valley or plateau. We focus on a generic and minimal model that includes only population subdivision into equivalent demes connected by global migration, and does not require significant size changes of the demes, environmental heterogeneity or specific geographic structure. We determine the optimal spee…
Using the continuous-time susceptible-infected-susceptible (SIS) model on networks, we investigate the problem of inferring the class of the underlying network when epidemic data is only available at population-level (i.e. the number of infected individuals at a finite set of discrete times of a single realisation of the epidemic), the only information likely to be available in real world settings. To tackle this, epidemics on networks are approximated by a Birth-and-Death process which keeps track of the number of infected nodes at population level. The rates of this surrogate model encode bo…
An SEIRS epidemic with disease fatalities is introduced in a growing population (modelled as a super-critical linear birth and death process). The study of the initial phase of the epidemic is stochastic, while the analysis of the major outbreaks is deterministic. Depending on the values of the parameters, the following scenarios are possible. i) The disease dies out quickly, only infecting few; ii) the epidemic takes off, the \textit{number} of infected individuals grows exponentially, but the \textit{fraction} of infected individuals remains negligible; iii) the epidemic takes off, the \text…
Mathematical disease modelling has long operated under the assumption that any one infectious disease is caused by one transmissible pathogen spreading among a population. This paradigm has been useful in simplifying the biological reality of epidemics and has allowed the modelling community to focus on the complexity of other factors such as population structure and interventions. However, there is an increasing amount of evidence that the strain diversity of pathogens, and their interplay with the host immune system, can play a large role in shaping the dynamics of epidemics. Here, we introd…
In this work, some new exact and approximate analytical solutions are obtained for the SIR epidemic model, which is formulated in terms of dimensionless variables and parameters. The susceptibles population (S) is in this way explicitly related to the infectives population (I) using the Lambert W function (both the principal and the secondary branches). A simple and accurate relation for the fraction of the population that does not catch the disease is also obtained. The explicit time dependences of the susceptibles, infectives and removed populations, as well as that of the epidemic curve are…
The death toll for Covid-19 may be reduced by dividing the population into two classes, the vulnerable and the fit, with different lockdown regimes. Instead of one reproduction number there now are four parameters. These make it possible to quantify the effect of the social distancing measures. There is a simple stochastic model for epidemics in a two type population. Apart from the size of the population of the vulnerable and the fit, and the initial number of infected in the two classes, only the four reproduction parameters are needed to run the two type Reed-Frost model. The program is sim…
By incorporating delayed epidemic information and self-restricted travel behavior into the SIS model, we have investigated the coupled effects of timely and accurate epidemic information and people's sensitivity to the epidemic information on contagion. In the population with only local random movement, whether the epidemic information is delayed or not has no effect on the spread of the epidemic. People's high sensitivity to the epidemic information leads to their risk aversion behavior and the spread of the epidemic is suppressed. In the population with only global person-to-person movement,…
We generalize the Susceptible-Infected-Removed model for epidemics to take into account generic effects of heterogeneity in the degree of susceptibility to infection in the population. We introduce a single new parameter corresponding to a power-law exponent of the susceptibility distribution that characterizes the population heterogeneity. We show that our generalized model is as simple as the original model which is contained as a limiting case. Because of this simplicity, numerical solutions can be generated easily and key properties of the epidemic wave can still be obtained exactly. In pa…
Motivated by recent epidemic outbreaks, including those of COVID-19, we solve the canonical problem of calculating the dynamics and likelihood of extensive outbreaks in a population within a large class of stochastic epidemic models with demographic noise, including the Susceptible-Infected-Recovered (SIR) model and its general extensions. In the limit of large populations, we compute the probability distribution for all extensive outbreaks, including those that entail unusually large or small (extreme) proportions of the population infected. Our approach reveals that, unlike other well-known …
Population heterogeneity, especially in individuals' contact networks, plays an important role in transmission dynamics of infectious diseases. For vaccine-preventable diseases, outstanding issues like vaccine hesitancy and availability of vaccines further lead to nonuniform coverage among groups, not to mention the efficacy of vaccines and the mixing pattern varying from one group to another. As the ongoing COVID-19 pandemic transitions to endemicity, it is of interest and significance to understand the impact of aforementioned population heterogeneity on the emergence and persistence of epid…
We study the impact of contact heterogeneity on epidemic dynamics. A system characterized by multiple susceptible populations is considered. The description of the spread of an infectious disease is obtained through the study of a system of Boltzmann-type equations for the number densities of social contacts of the introduced compartments. A macroscopic system of equations characterizing observable effects of the epidemic is then derived to assess the impact of contact heterogeneity.
Investigations of a possible connection between population density and the propagation and magnitude of epidemics have so far led to mixed and unconvincing results. There are three reasons for that. (i) Previous studies did not focus on the appropriate density interval. (ii) For the density to be a meaningful variable the population must be distributed as uniformly as possible. If an area has towns and cities where a majority of the population is concentrated its average density is meaningless. (iii) In the propagation of an epidemic the initial proportion of susceptibles (that is to say perso…
Compartmental equations are primary tools in disease spreading studies. Their predictions are accurate for large populations but disagree with empirical and simulated data for finite populations, where uncertainties become a relevant factor. Starting from the agent-based approach, we investigate the role of uncertainties and autocorrelation functions in SIS epidemic model, including their relationship with epidemiological variables. We find new differential equations that take uncertainties into account. The findings provide improved predictions to the SIS model and it can offer new insights f…
Empirical studies show that epidemiological models based on an epidemic's initial spread rate often fail to predict the true scale of that epidemic. Most epidemics with a rapid early rise die out before affecting a significant fraction of the population, whereas the early pace of some pandemics is rather modest. Recent models suggest that this could be due to the heterogeneity of the target population's susceptibility. We study a computer malware ecosystem exhibiting spread mechanisms resembling those of biological systems while offering details unavailable for human epidemics. Rather than com…
The outbreak of an infectious disease in a human population can lead to individuals responding with preventive measures in an attempt to avoid getting infected. This leads to changes in contact patterns. However, as we show in this paper, rational behaviour at the individual level, such as social distancing from infectious contacts, may not always be beneficial for the population as a whole. We use epidemic network models to demonstrate the potential negative consequences at the population level. We take into account the social structure of the population through several network models. As the…
The outbreak of COVID-19, beginning in 2019 and continuing through the time of writing, has led to renewed interest in the mathematical modeling of infectious disease. Recent works have focused on partial differential equation (PDE) models, particularly reaction-diffusion models, able to describe the progression of an epidemic in both space and time. These studies have shown generally promising results in describing and predicting COVID-19 progression. However, people often travel long distances in short periods of time, leading to nonlocal transmission of the disease. Such contagion dynamics …
In this paper we consider an SIRS epidemic model under a general assumption of density-dependent mortality. We prove the global stability of the disease-free equilibrium and propose a Lyapunov function that allows to demonstrate the global stability of the (unique) endemic state under broad conditions.
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-popul…
The ability to directly record human face-to-face interactions increasingly enables the development of detailed data-driven models for the spread of directly transmitted infectious diseases at the scale of individuals. Complete coverage of the contacts occurring in a population is however generally unattainable, due for instance to limited participation rates or experimental constraints in spatial coverage. Here, we study the impact of spatially constrained sampling on our ability to estimate the epidemic risk in a population using such detailed data-driven models. The epidemic risk is quantif…
Most infectious diseases including more than half of known human pathogens are not restricted to just one host, yet much of the mathematical modeling of infections has been limited to a single species. We investigate consequences of a single epidemic propagating in multiple species and compare and contrast it with the endemic steady state of the disease. We use the two-species Susceptible-Infected-Recovered (SIR) model to calculate the severity of post-epidemic collapses in populations of two host species as a function of their initial population sizes, the times individuals remain infectious,…
A new method is proposed to infer unobserved epidemic sub-populations by exploiting the synchronization properties of multistrain epidemic models. A model for dengue fever is driven by simulated data from secondary infective populations. Primary infective populations in the driven system synchronize to the correct values from the driver system. Most hospital cases of dengue are secondary infections, so this method provides a way to deduce unobserved primary infection levels. We derive center manifold equations that relate the driven system to the driver system and thus motivate the use of sync…
This manuscript proposes a model of $n$ distinct populations interaction structured SEIR to describe the spread of COVID-19 pandemic diseases. The proposed model has the flexibility to include geographically separated communities as well as taking into account aging population groups and their interactions. We show that distinct assumptions on the dynamics of the proposed model lead to a plateau-like curve of the infected population, reflecting collected data from large countries like Brazil. Such observations aim to the following conjecture: " Covid-19 diseased diffusion from the capitals to …
Developing robust, quantitative methods to optimize resource allocations in response to epidemics has the potential to save lives and minimize health care costs. In this paper, we develop and apply a computationally efficient algorithm that enables us to calculate the complete probability distribution for the final epidemic size in a stochastic Susceptible-Infected-Recovered (SIR) model. Based on these results, we determine the optimal allocations of a limited quantity of vaccine between two non-interacting populations. We compare the stochastic solution to results obtained for the traditional…
University of Oxford（牛津大学）；Massachusetts Institute of Technology (MIT)（麻省理工学院）；TU Delft（代尔夫特理工大学）；German Aerospace Center (DLR)（德国航空航天中心）；Proxima Robotics srl（普罗xima机器人有限公司）
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The SARS-CoV-2 pandemic has caused significant mortality and morbidity worldwide, sparing almost no community. As the disease will likely remain a threat for years to come, an understanding of the precise influences of human demographics and settlement, as well as the dynamic factors of climate, susceptible depletion, and intervention, on the spread of localized epidemics will be vital for mounting an effective response. We consider the entire set of local epidemics in the United States; a broad selection of demographic, population density, and climate factors; and local mobility data, trackin…
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We study contact epidemic models for the spread of infective diseases in finite populations. The size dependence enters in the infection rate. The dynamics of such models is then analyzed within the deterministic approximation, as well as in terms of a stochastic formulation. At the level of the deterministic equations, we deduce relations between the parameters of the model for the disease to become endemic. Within the stochastic formulation, it is possible to write recursion relations for the probability distribution, that lead to exact expression for some of its moments and check the validi…
To forecast the time dynamics of an epidemic, we propose a discrete stochastic model that unifies and generalizes previous approaches to the subject. Viewing a given population of individuals or groups of individuals with given health state attributes as living in and moving between the nodes of a graph, we use Monte-Carlo Markov Chain techniques to simulate the movements and health state changes of the individuals according to given probabilities of stay that have been preassigned to each of the nodes. We utilize this model to either capture and predict the future geographic evolution of an e…
We investigate a time-dependent spatial vector-host epidemic model with non-coincident domains for the vector and host populations. The host population resides in small non-overlapping sub-regions, while the vector population resides throughout a much larger region. The dynamics of the populations are modeled by a reaction-diffusion-advection compartmental system of partial differential equations. The disease is transmitted through vector and host populations in criss-cross fashion. We establish global well-posedness and uniform a prior bounds as well as the long-term behavior. The model is ap…
A simple, but ``classical``, stochastic model for epidemic spread in a finite, but large, population is studied. The progress of the epidemic can be divided into three different phases that requires different tools to analyse. Initially the process is approximated by a branching process. It is discussed for how long time this approximation is valid. When a non-negligible proportion of the population is already infected the process can be studied using differential equations. In a final phase the spread will fade out. The results are used to investigate what happens if two strains of infectious…
In this paper, we provide a straightforward approach to defining and deriving the key epidemiological quantity, the basic reproduction number, $R_0$, for Markovian epidemics in structured populations. The methodology derived is applicable to, and demonstrated on, both $SIR$ and $SIS$ epidemics and allows for population as well as epidemic dynamics. The approach taken is to consider the epidemic process as a multitype process by identifying and classifying the different types of infectious units along with the infections from, and the transitions between, infectious units. For the household mod…
We study how international flights can facilitate the spread of an epidemic to a worldwide scale. We combine an infrastructure network of flight connections with a population density dataset to derive the mobility network, and then we define an epidemic framework to model the spread of the disease. Our approach combines a compartmental SEIRS model with a graph diffusion model to capture the clusteredness of the distribution of the population. The resulting model is characterised by the dynamics of a metapopulation SEIRS, with amplification or reduction of the infection rate which is determined…
We predict the future course of ongoing susceptible-infected-susceptible (SIS) epidemics on regular, Erdős-Rényi and Barabási-Albert networks. It is known that the contact network influences the spread of an epidemic within a population. Therefore, observations of an epidemic, in this case at the population-level, contain information about the underlying network. This information, in turn, is useful for predicting the future course of an ongoing epidemic. To exploit this in a prediction framework, the exact high-dimensional stochastic model of an SIS epidemic on a network is approximated by a …
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