In the recent COVID-19 pandemic we assisted at a sequence of epidemic waves intertwined by anomalous fade-outs with periods of low but persistent epidemic prevalence. These long-living epidemic states complicate epidemic control and challenge current modeling approaches as classical epidemic models fail to explain their emergence. Inspired by this phenomenon, we propose a simple mechanism able to reproduce several features observed in real data. Specifically, here we introduce a modification of the Susceptible-Infected-Recovered (SIR) model in a meta-population framework where a small inflow of infected individuals accounts for undetected internal or imported cases. Focusing on a regime where this external seeding is so small that cannot be detected from the analysis of epidemic curves, we find that outbreaks of finite duration percolate in time resulting in overall low but long-living epidemic states below and above the epidemic threshold. Using a two-state description of the local dynamics, we can extract analytical predictions for the phase space. The comparison with epidemic data demonstrates that our model is able to reproduce some critical signatures observed in COVID-19 spreading in England. Finally, our findings defy our understanding of the concept of epidemic threshold and its relationship with outbreaks survival for disease control.
Most countries are suffering severely from the ongoing covid-19 pandemic despite various levels of preventive measures. A common question is if and when a country or region will reach herd immunity $h$. The classical herd immunity level $h_C$ is defined as $h_C=1-1/R_0$, where $R_0$ is the basic reproduction number, for covid-19 estimated to lie somewhere in the range 2.2-3.5 depending on country and region. It is shown here that the disease-induced herd immunity level $h_D$, after an outbreak has taken place in a country/region with a set of preventive measures put in place, is actually subst…
Here we show that herd immunity is not a measure we can implement, trust, control or depend on, and that waiting for herd immunity to play a role is tantamount to hope for the best. Herd immunity is not some protection, as it has been referred to, but a reduction in the chance of infection that a population can achieve only if more infections occur, which is paradoxical. The discussion on when there will be enough herd immunity for the COVID-19 epidemics to halt, delivers the public and policy makers the wrong message that we can wait for something to happen to relax containment measures. Here…
We study a SEIR model considered by Gomes et al. \cite{Gomes2020} and Aguas et al. \cite{Aguas2020} where different individuals are assumed to have different levels of susceptibility or exposure to infection. Under this heterogeneity assumption, epidemic growth is effectively suppressed when the percentage of population having acquired immunity surpasses a critical level - the herd immunity threshold - that is lower than in homogeneous populations. We find explicit formulas to calculate herd immunity thresholds and stable configuration, and explore extensions of the model.
A quantitative COVID-19 model that incorporates hidden asymptomatic patients is developed, and an analytic solution in parametric form is given. The model incorporates the impact of lockdown and resulting spatial migration of population due to announcement of lockdown. A method is presented for estimating the model parameters from real-world data. It is shown that increase of infections slows down and herd immunity is achieved when symptomatic patients are 4-6\% of the population for the European countries we studied, when the total infected fraction is between 50-56 \%. Finally, a method for …
We performed a thorough sensitivity analysis of the herd immunity threshold for discrete-time SIR compartmental models with a static network structure. We find unexpectedly that these models violate classical intuition which holds that the herd immunity threshold should monotonically increase with the transmission parameter. We find the existence of bifurcations in the herd immunity threshold in the high transmission probability regime. The extent of these bifurcations is modulated by the graph heterogeneity, the recovery parameter, and the network size. In the limit of large, well-mixed netwo…
We model and calculate the fraction of infected population necessary to reach herd immunity, taking into account the heterogeneity in infectiousness and susceptibility, as well as the correlation between those two parameters. We show that these cause the effective reproduction number to decrease more rapidly, and consequently have a drastic effect on the estimate of the necessary percentage of the population that has to contract the disease for herd immunity to be reached. We quantify the difference between the size of the infected population when the effective reproduction number decreases be…
The contact structure of a population plays an important role in transmission of infection. Many ``structured models'' capture aspects of the contact structure through an underlying network or a mixing matrix. An important observation in such models, is that once a fraction $1-1/\mathcal{R}_0$ has been infected, the residual susceptible population can no longer sustain an epidemic. A recent observation of some structured models is that this threshold can be crossed with a smaller fraction of infected individuals, because the disease acts like a targeted vaccine, preferentially immunizing highe…
In constant parameter compartmental models an early onset of herd immunity is at odds with estimates of R values from early stage growth. This paper utilizes a result from the theory of interest rate modeling, namely a bond pricing formula of Vasicek, and an approach inspired by a foundational result in statistics, de Finetti's Theorem, to show how the modeling discrepancy can be explained. Moreover the difference between predictions of classic constant parameter epidemiological models and those with variation and stochastic evolution can be reduced to simple "convexity" formulas. A novel feat…
COVID-19 will be a continuous threat to human population despite having a few vaccines at hand until we reach the endemic state through natural herd immunity and total immunization through universal vaccination. However, the vaccine acts as a practical tool for reducing the massive public health problem and the emerging economic consequences that the continuing COVID -19 epidemic is causing worldwide, while the vaccine efficacy wanes. In this work, we propose and analyze an epidemic model of Susceptible-Exposed-Infected-Recovered-Vaccinated (SEIRV) population taking into account the rate of va…
We analyse an extended SIRS epidemic model in which immunity at the individual level wanes gradually at exponential rate, but where the waning rate may differ between individuals, for instance as an effect of differences in immune systems. The model also includes vaccination schemes aimed to reach and maintain herd immunity. We consider both the informed situation where the individual waning parameters are known, thus allowing selection of vaccinees being based on both time since last vaccination as well as on the individual waning rate, and the more likely uninformed situation where individua…
Epidemics generally spread through a succession of waves that reflect factors on multiple timescales. On short timescales, super-spreading events lead to burstiness and overdispersion, while long-term persistent heterogeneity in susceptibility is expected to lead to a reduction in the infection peak and the herd immunity threshold (HIT). Here, we develop a general approach to encompass both timescales, including time variations in individual social activity, and demonstrate how to incorporate them phenomenologically into a wide class of epidemiological models through parameterization. We deriv…
Countries highly exposed to incoming traffic from China were expected to be at the highest risk of COVID-19 spread. However, COVID-19 case numbers (infection levels) are negatively correlated with incoming traffic-level. Moreover, infection levels are positively correlated with population-size, while the latter should only affect infection-level once herd immunity is reached. These could be explained if a low-virulence strain (LVS) began spreading a few months earlier from China, providing immunity from the later emerging known SARS-CoV-2 high-virulence strain (HVS). We find that the dynamics …
HUN-REN Alfréd Rényi Institute of Mathematics（匈牙利科学院阿尔弗雷德·雷尼数学研究所）；Faculty of Electrical Engineering, Mathematics and Computer Science, Delft University of Technology（代尔夫特理工大学电气、数学和计算机科学学院）；Institute for Hygiene and Applied Immunology, Medical University of Vienna（维也纳医科大学卫生与应用免疫学研究所）；Department of Network and Data Science, Central European University（中欧大学网络与数据科学系）
Until a vaccine or therapy is found against the SARS-CoV-2 coronavirus, reaching herd immunity appears to be the only mid-term option. However, if the number of infected individuals decreases and eventually fades only beyond this threshold, a significant proportion of susceptible may still be infected until the epidemic is over. A containment strategy is likely the best policy in the worst case where no vaccine or therapy is found. In order to keep the number of newly infected persons to a minimum, a possible strategy is to apply strict containment measures, so that the number of susceptible i…
Genotype networks drive oscillating endemicity and epidemic trajectories in viral evolution
University of Zaragoza（萨拉戈萨大学）；Institute of Biocomputation and Physics of Complex Systems (BIFI), University of Zaragoza（萨拉戈萨大学复杂系统生物计算与物理研究所 (BIFI)）；Universitat Rovira i Virgili（罗维拉-伊-维尔吉利大学）；Pacific Northwest National Laboratory（太平洋西北国家实验室）；University of Kobe（神户大学）
The current COVID-19 pandemic and subsequent lockdowns have highlighted the close and delicate relationship between a country's public health and economic health. Macroeconomic models that use preexisting epidemic models to calculate the impacts of a disease outbreak are therefore extremely useful for policymakers seeking to evaluate the best course of action in such a crisis. We develop an SIR model of the COVID-19 pandemic that explicitly considers herd immunity, behavior-dependent transmission rates, remote workers, and indirect externalities of lockdown. This model is presented as an exit …
This paper introduces a new multi-strain epidemic model with reinfection and cross-immunity to provide insights into the resurgence of the COVID-19 epidemic in an area with reportedly high seroprevalence due to a largely unmitigated outbreak: the state of Amazonas, Brazil. Although high seroprevalence could have been expected to trigger herd immunity and prevent further waves in the state, we have observed persistent levels of infection after the first wave and eventually the emergence of a second viral strain just before an augmented second wave. Our experiments suggest that the persistent le…
We consider the $SEIRS$ epidemiology model with such features of the COVID-19 outbreak as: abundance of unidentified infected individuals, limited time of immunity and a possibility of vaccination. Within a compartmental realization of this model, we found the disease-free and the endemic stationary states. They exist in their respective restricted regions of the le via linear stability analysis. The expression for the basic reproductive number is obtained as well. The positions and heights of a first peak for the fractions of infected individuals are obtained numerically and are fitted to sim…
Bareilly College（巴雷利学院）；CSIR-Central Electronics Engineering Research Institute（CSIR-中央电子工程研究所）；Jawaharlal Nehru University（贾瓦哈拉尔·尼赫鲁大学）
To combat the coronavirus disease 2019 (COVID-19) pandemic, the world has vaccination, plasma therapy, herd immunity, and epidemiological interventions as few possible options. The COVID-19 vaccine development is underway and it may take a significant amount of time to develop the vaccine and after development, it will take time to vaccinate the entire population, and plasma therapy has some limitations. Herd immunity can be a plausible option to fight COVID-19 for small countries. But for a country with huge population like India, herd immunity is not a plausible option, because to acquire he…
Using a simple economic model in which social-distancing reduces contagion, we study the implications of waning immunity for the epidemiological dynamics and social activity. If immunity wanes, we find that COVID-19 likely becomes endemic and that social-distancing is here to stay until the discovery of a vaccine or cure. But waning immunity does not necessarily change optimal actions on the onset of the pandemic. Decentralized equilibria are virtually independent of waning immunity until close to peak infections. For centralized equilibria, the relevance of waning immunity decreases in the pr…
The COVID-19 pandemic has hit different parts of the world differently: some regions are still in the rise of the first wave, other regions are now facing a decline after a first wave, and yet other regions have started to see a second wave. The current immunity level $\hat i$ in a region is closely related to the cumulative fraction infected, which primarily depends on two factors: a) the initial potential for COVID-19 in the region (often quantified by the basic reproduction number $R_0$), and b) the timing, amount and effectiveness of preventive measures put in place. By means of a mathemat…
I employ a simple mathematical model of an epidemic process to evaluate how four basic quantities: the reproduction number (R), the numbers of sensitive (S) and infectious individuals(I), and total community size (N) affect strategies to control COVID-19. Numerical simulations show that strict suppression measures at the beginning of an epidemic can create low infectious numbers, which thereafter can be managed by mitigation measures over longer periods to flatten the epidemic curve. The stronger the suppression measure, the faster it achieves the low numbers of infections that are conducive t…
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…
Mathematical modelling of the spread of epidemics has been an interesting challenge in the field of epidemiology. The SIR Model proposed by Kermack and McKendrick in 1927 is a prototypical model of epidemiology. However, it has its limitations. In this paper, we show two independent ways of generalizing this model, the first one if the vaccine isn't discovered or ready to use and the next one, if the vaccine is discovered and ready to use. In the first part, we have pointed out a major over-simplification, i.e., assumption of variation of the time derivatives of the variables with the linear o…
In this paper we conduct a simulation study of the spread of an epidemic like COVID-19 with temporary immunity on finite spatial and non-spatial network models. In particular, we assume that an epidemic spreads stochastically on a scale-free network and that each infected individual in the network gains a temporary immunity after its infectious period is over. After the temporary immunity period is over, the individual becomes susceptible to the virus again. When the underlying contact network is embedded in Euclidean geometry, we model three different intervention strategies that aim to contr…
As infectious disease outbreaks emerge, public health agencies often enact vaccination and social distancing measures to slow transmission. Their success depends on not only strategies and resources, but also public adherence. Individual willingness to take precautions may be influenced by global factors, such as news media, or local factors, such as infected family members or friends. Here, we compare three modes of epidemiological decision-making in the midst of a growing outbreak. Individuals decide whether to adopt a recommended intervention based on overall disease prevalence, the proport…
We study variants of the SEIR model for interpreting some qualitative features of the statistics of the Covid-19 epidemic in France. Standard SEIR models distinguish essentially two regimes: either the disease is controlled and the number of infected people rapidly decreases, or the disease spreads and contaminates a significant fraction of the population until herd immunity is achieved. After lockdown, at first sight it seems that social distancing is not enough to control the outbreak. We discuss here a possible explanation, namely that the lockdown is creating social heterogeneity: even if …
In this paper, we provide insights on how much testing and social distancing is required to control COVID-19. To this end, we develop a compartmental model that accounts for key aspects of the disease: 1) incubation time, 2) age-dependent symptom severity, and 3) testing and hospitalization delays; the model's parameters are chosen based on medical evidence, and, for concreteness, adapted to the German situation. Then, optimal mass-testing and age-dependent social-distancing policies are determined by solving optimal control problems both in open loop and within a model predictive control fram…
After the first lockdown in response to the COVID-19 outbreak, many countries faced difficulties in balancing infection control with economics. Due to limited prior knowledge, economists began researching this issue using cost-benefit analysis and found that infection control processes significantly affect economic efficiency. A UK study used economic parameters to numerically demonstrate an optimal balance in the process, including keeping the infected population stationary. However, universally applicable knowledge, which is indispensable for the guiding principles of infection control, has …
