Abstract
Social distancing as one of the main non-pharmaceutical interventions can help slow down the spread of diseases, like in the COVID-19 pandemic. Effective social distancing, unless enforced as drastic lockdowns and mandatory cordon sanitaire, requires consistent strict collective adherence. However, it remains unknown what the determinants for the resultant compliance of social distancing and their impact on disease mitigation are. Here, we incorporate into the epidemiological process with an evolutionary game theory model that governs the evolution of social distancing behaviour. In our model, we assume an individual acts in their best interest and their decisions are driven by adaptive social learning of the real-time risk of infection in comparison with the cost of social distancing. We find interesting oscillatory dynamics of social distancing accompanied with waves of infection. Moreover, the oscillatory dynamics are dampened with a non-trivial dependence on model parameters governing decision-makings and gradually cease when the cumulative infections exceed the herd immunity. Compared to the scenario without social distancing, we quantify the degree to which social distancing mitigates the epidemic and its dependence on individuals’ responsiveness and rationality in their behaviour changes. Our work offers new insights into leveraging human behaviour in support of pandemic response.
Keywords: behavioural epidemiology, evolutionary game theory, disease dynamics
1. Introduction
Emerging novel zoonotic diseases, such as Zika [1], Ebola [2] and the most recently COVID-19 [3], have imposed great threats to global health and humanity [4]. Some of these new diseases are caused by respiratory viruses and highly contagious through proximity transmissions, and may turn into an unprecedented pandemic well before effective treatments and vaccines have been developed and widely deployed. In this case, the world may have to resort to non-pharmaceutical interventions (NPI), such as face covering and social distancing so as to mitigate disease impact before effective pharmaceutical interventions become available. However, the ultimate effectiveness of NPI measures is highly contingent on compliance and adherence, since NPI is usually not a one-off measure, but rather requires repeated, consistent adherence in order to reduce potential transmission routes of contracting the infection. From this perspective, human behaviour plays an important role in impacting the course of a pandemic outbreak as well as the health outcome.
In recent years, there has been growing interest in understanding social factors in epidemiology (see, for example, [5] for a brief review). In the field of behavioural epidemiology, of particular interest is the use of disease-behaviour interaction models for this purpose [6]. Prior work has extensively used this framework to study how vaccine compliance can be influenced by a wide range of factors [7–9], ranging from vaccine scares [10] to disease awareness [11]. The feedback loop between behavioural change and disease prevalence gives rise to a variety of interesting, non-trivial dynamics [12–15], e.g. the hysteresis effect [16]. Among others, an important approach is combining evolutionary game theory with epidemiological models [7,17,18]. Evolutionary game theory provides a general mathematical framework for modelling behavioural changes in a population driven by both social influence and self-interest. In the past decades, the approach of replicator dynamics has been commonly used to model social learning/imitation process, and particularly the spread of behaviour (social contagion), in a range of important real-world problems [19], from peer punishment [20] over cooperation [21], altruistic punishment [22], honesty [23], trust [24] and moral behaviour in general [25] to antibiotic usage [26].
Unlike vaccination, social distancing effort of an individual requires repeated decisions whether or not to comply by evaluating the necessity of doing so throughout the epidemic, despite public health recommendations or even mandates [27]. The cost of social distancing is not negligible, but rather has a huge impact on the economic status and well-being of people [28]. Previous work has modelled social distancing as a differential game [29], that is, individuals try to maximize their payoffs by adjusting their effort in social distancing (namely, the level of exposure to potential transmission routes) by comparing the risk of contracting the disease with the cost of social distancing. Their numerical results show that the collective dynamics of social distancing would approach to a steady level (i.e. a Nash equilibrium with constant effort for social distancing) without any oscillatory dynamics [29]. While this prior study sheds useful insights for social distancing from the game theory perspective, it remains largely unknown how the rationality and the responsiveness of individuals in reacting to an epidemic would impact the compliance level of social distancing.
Social distancing is costly, yet if not optimized for timing and duration and intensity, it would lead to wasted effort [30,31]. Combined with real data, the impact of social distancing can also be quantitatively assessed and optimized for past pandemics like influenza [32,33]. Noteworthy, there have been efforts to predict and quantify the effectiveness of reactive distancing on the COVID-19 pandemic, in anticipation of multiple waves of infections in the coming years [34,35].
Aside from individual perspective, the optimization of disease control is often studied using optimal control theory by assuming a central social planner aiming to minimize the cost of disease outbreak [36–39]. While these results are insightful from the perspective of population optima [40] (that is, optimized policies are complied uniformly in the population), it is challenging to attain these goals in practice due to compliance issues.
To shed light on driving factors of compliance levels of social distancing, here we take into account important aspects of human decision-making—bounded rationality [41] and loss aversion [42]—which is informed by the real-time disease prevalence, and prompted by peers’ choice. We incorporate into the epidemiological process with an evolutionary game dynamics of social distancing behaviour. Individuals decide on whether or not to commit to social distancing by weighing the risk of infection with the cost of social distancing. The responsiveness parameter in our model modulates the relative time scale of individuals revisiting their social distancing decisions, as compared to the pace of an unfolding epidemic. We introduce bounded rationality that individuals are not necessarily using the best response but rather with some probability of changing their behaviour.
In this work, we find an interesting oscillatory tragedy of the commons in the collective dynamics of social distancing. Individuals are inclined to social distancing when the disease prevalence is above a threshold that depends on the transmissibility of the disease and the relative cost of social distancing versus contracting the disease. As the epidemic curve is being flattened, individuals consequently feel more safe not to practise social distancing, thereby causing the decline in the compliance of social distancing and further resulting in a resurgence of disease outbreaks in the population. Even though such reactive social distancing is hardly able to help reach the optimality of disease mitigation, it can avoid the overshooting of infected individuals which typically happens in an susceptible–infected–recovered (SIR) model in the absence of any interventions. We also find non-trivial dependence of the effectiveness of social distancing, measured by the fraction of susceptible individuals who would become infected without social distancing, on model parameters governing individuals’ rationality and responsiveness.
2. Model and methods
(a). Model
Our model is basically a combination of the classical SIR model with the replicator equation: In a well-mixed infinite population each individual is either susceptible, infected or recovered. Moreover, each susceptible individual can at each time choose to either practise social distancing or not to practise social distancing. If the individual practises social distancing they cannot become infected. If they do not practise social distancing they become infected in an encounter with an infected individual with probability β > 0. At each time an infected recovers with probability γ > 0 (figure 1).
We denote the proportion of susceptible individuals at time t by S(t), the proportion of infected by I(t), and the proportion of removed by R(t). Furthermore, we denote by the proportion of susceptible individuals that practise social distancing. We denote the initial conditions by I0 = I(0), S0 = S(0) as well as . A susceptible individual determines his strategy based on a cost-benefit analysis. Hence, by πsd we denote the payoff of social distancing, and by πnsd the payoff of no social distancing. In our model, the perceived cost of social distancing is Csd > 0 at each time t. Thus, we have
πnsd depends on two factors: the perceived cost of infection that we denote by CI > 0 and the risk of infection. The risk of infection in time (t, t + 1) without social distancing is given by
Therefore, the payoff of not socially distancing is given by
Hence, the dynamics of our model are given by the following system of ordinary differential equations (ODEs):
2.1 |
Here, ω is a responsiveness parameter, determining the time scale for updating the social distancing behaviour. κ is a rationality parameter. For large κ individuals change their strategy if the payoff of the other strategy is larger. For small κ only a fraction of the susceptible individuals depending on the difference in payoff change their strategy. The behaviour of this model is illustrated in figure 2 for different parameters. For this figure as well as for all other figures, we used the Matlab method ode23, which is an implementation of the Bogacki–Shampine method—an explicit Runge–Kutta (2,3) pair.
(b). Perfect adaption
In Model (2.1), the dynamics of social distancing change to direct the amount of infected I towards the amount where πsd = πnsd, i.e. towards . Assuming that this adaption works perfectly, we obtain the following model given by the ODEs.
This model is given by the ODEs
2.2 |
with S(t) = 1 − IPA(t) − RPA(t), , and with initial condition
An illustration of this model is given in figure 3.
Then, the total amount of people that get infected RPA(∞) is given by
where W denotes the Lambert W function. Thus, in the case of perfect adaption, we can achieve
by choosing Cd/CI → 0. We want to use this model of perfect adaption to understand how the total amount of infected R(∞) in Model (2.1) depends on the parameters Csd, CI, ω, κ.
3. Results
(a). Oscillatory tragedy of the commons
In Model (2.1), the cost of social distancing and no social distancing are equal at time t if
If , then is increasing. If then is decreasing. On the other hand, if is sufficiently large, this causes a decrease in I and if is sufficiently small this causes an increase in I. If the amount of infections is high, people are more aware of the disease and practise social distancing. As soon as the amount of infections is small again, this awareness fades and people do not feel the need to practise social distancing anymore. As a result, more people become infected again leading to a higher awareness and more people practising social distancing. We refer to this feedback loop as oscillatory tragedy of the commons. Instead of high compliance to social distancing until the disease has died of, we find a decrease in individuals practising social distancing when the amount of infected is sufficiently small. This then causes another rise of infections. We can observe this in Model (2.1) as I oscillates around with decreasing amplitude until the peak of the oscillations is smaller than (figure 4).
(b). Social distancing saves lives
When comparing Model (2.1) to the SIR model, we immediately note that the total number of infections can be significantly smaller with social distancing (figure 5). Essentially, this means that voluntary social distancing can significantly reduce the total amount of infections R(∞). However, we also note that infections after the first wave of infection only emerge due to the oscillatory tragedy of the commons. If social distancing was practised until I = 0, we would have a much smaller R(∞).
Perfect adaption and Model (2.1) significantly reduce the total amount of infections compared to the SIR model. This is especially apparent for small and slow adaption, i. e. small ω and κ. This is illustrated in figure 6.
Essentially, the explanation for this behaviour relates to herd immunity. Social distancing flattens the curve. Instead of one large wave of infections as in the SIR model, in Model (2.1) we can have several waves of infection with smaller peaks. An example of this is illustrated in figure 5. In the SIR model, herd immunity occurs if
Thus, I is increasing until S = γ/β and then is monotonically decreasing. Even though, we have achieved some kind of herd immunity at this point, the high number of infected I still causes a high amount of new infections after herd immunity. Thus, the total amount of infections R(∞) is significantly larger than needed to obtain herd immunity. In Model (2.1), the dynamics are much more complicated. However, what remains as in the SIR model, is that as soon as (or at latest at this point) S < γ/β the amount of infected I is monotonically decreasing, since then we have
We denote the amount of infected when herd immunity is obtained by IHI. With social distancing IHI can become significantly smaller since social distancing significantly reduces the amount of infection. Other factors influencing the amount of new infection after herd immunity are the amount of recovered when herd immunity is achieved (denoted by RHI) as well as the amount of people practising social distancing. RHI can be much larger when social distancing is practised due to the spread of infections over a longer time period. People that practise social distancing further reduce the amount of new infections.
Together, all these factors cause a significant decrease in new infections after herd immunity is achieved. Since small IHI mostly coincides with large RHI as well as high , we focus on IHI here. When choosing the parameters ω, κ, Csd, CI such that IHI is small, RHI and many people practice social distancing, we can even achieve R(∞) to be near the herd immunity threshold 1 − γ/β. For an illustration of this, see figure 7.
This also explains why perfect adaption causes larger total amounts of infections than Model (2.1). For perfect adaption, herd immunity is obtained for with while IHI often is significantly smaller in Model (2.1). In particular, if we have while S is close to γ/β, small increases in cause a decrease in I. Hence, in most cases. For instance, in figure 7a, we have while IHI can be much smaller in Model (2.1). However, we once again remember the oscillatory tragedy of the commons, i.e. that higher compliance to social distancing when I is small could lead to much smaller R(∞).
Next, we want to analyse how the perceived cost of social distancing Csd as well as the perceived cost of infection CI influence the total amount of infections R(∞).
(c). Larger cost of infection and smaller cost of social distancing reduce infections
As one might expect, if the cost of infection CI increases or the cost of social distancing Csd decreases, this induces an increase in the amount of people practising social distancing and thereby a decrease in infections. This behaviour becomes quite apparent in figure 8.
We can observe a similar tendency for CI. Though, here we have larger oscillations in the total size of infections. These oscillations decrease in their amplitude and level off at RPA(∞). An example for this behaviour can be seen in figure 9.
An explanation for this behaviour is connected to the observation made in §b that is illustrated in figure 7. To reduce R(∞), (among other factors) IHI has to be small. One way to achieve this is to reduce , the threshold that I oscillates around. Therefore, smaller Csd as well as larger CI tend to cause a decrease in R(∞). However, this does not yet explain how the oscillations occur. For this purpose, we have a look atfigure 10. Here, we see that reducing has two opposing effects on R(∞):
-
(1)
A decrease in causes a decrease in the size of the waves of infections and thus a decrease in IHI. This causes a decrease in R(∞).
-
(2)
When decreasing too much, this can lead to the development of a new wave of infections. When this occurs, we have an increase in I, before herd immunity is obtained. This causes a larger IHI. Therefore, we have an increase in the total amount of infections R(∞) when a new wave of infections develops.
This leads to the oscillations, that we observed in figure 9. When reducing Csd we first see a decrease in R(∞) (caused by smaller waves of infection and a smaller IHI) followed by an increase (induced by a new wave of infections that leads to an increase in IHI).
(d). Faster responses and higher rationality increase infections
Two other important factors determining R(∞) are the responsiveness ω and the rationality parameter κ. In Model (2.1), a larger ω causes faster adaption of social distancing to the amount of infected. This has two opposing effects.
-
(1)
On the one hand, faster adaptions causes a decrease in the duration of the waves of infection with smaller maxima and larger minima.
-
(2)
On the other hand, these smaller waves of infection can cause the development of another wave of infection. In particular, if herd immunity is obtained before this new wave recedes, this leads to an increase in IHI. Thus, leading to an increase of infections.
Overall, we thus have oscillations in R(∞) depending on ω. An example of this is illustrated in figure 11.
With increasing ω the deviations of I from are decreasing due to faster adaption. This leads to a decrease in the amplitude of the oscillations in R(∞) and to R(∞) levelling off approximately at RPA(∞). As explained before, we mostly have . Therefore, larger deviations from perfect adaption where cause a decrease in R(∞). An example of this behaviour is illustrated in figure 12.
The rationality parameter κ has a nearly similar effect as ω on the dynamics of our model. In Model (2.1), a large rationality parameter κ means that individuals change their strategy as soon as the payoff of infection becomes larger than the payoff of social distancing and vice versa. Therefore, large κ induce faster adaption of and therefore smaller oscillations of I around . An increase in κ thus causes a decrease in the duration in the waves of infection as well as smaller maxima and larger minima. Hence, a change in κ has a similar effect on R(∞) as a change in ω. Here as well, we have oscillations caused by the development of new waves of infection, that are decreasing and levelling off at RPA(∞). Thus, R(∞) tends to decrease for smaller κ (figure 13).
4. Discussion and conclusion
Social distancing is often used in combination with other control measures such as mask wearing, and testing and isolation. It is worthy of further investigation to account for individual preferences in their adoption choices when multiple interventions for disease mitigation are available. Generally speaking, individuals become less vigilant and feel less need to follow disease intervention measures suggested by public health officials, if the epidemic curve is being bent down, but as a result, the uptick of cases in turn causes individuals to increase their compliance levels. The feedback loop of this sort gives rise to an oscillatory dynamics of behavioural compliance and disease prevalence, as reported in the present work. Similar phenomena have previously been studied in the context of eco-evolutionary dynamics where the payoff structure of individual interactions can be regulated by the environmental feedback [43–45].
Social distancing can be regarded as an altruistic behaviour that incurs a cost to oneself but collectively benefits other community members especially these vulnerable in the population. Thus monetary or non-monetary means can be used to incentivize non-compulsory social distancing. For example, during the COVID-19 health crisis, governments have subsidized the cost of staying at home through tax reduction or other stimulus packages for both workers and their employers [28]. Besides, an individual who opts for social distancing can create a positive psychological reward, which in fact reduces the perceived overall cost of social distancing. As shown in several experimental works [46–48], encouraging altruistic social distancing, especially if people can afford to do so, through promoting a strong sense of community, empathy and compassion [49], can lead to desired compliance of social distancing. In this sense, promoting human cooperation in the social dilemma of disease control is a new promising direction for future work.
While our proof-of-principle model offers enlightening insights into understanding compliance issues in the dilemma of social distancing, targeted social distancing can be investigated by further accounting for individual heterogeneity as the attack rate and mortality rate of infectious diseases, such as the influenza [50,51] and the COVID-19 pandemic [52], are age-dependent. Thus, extending our model with an age structure will be useful to quantify the heterogeneity in both the risk of infections and the cost of social distancing for each age group. This consideration parameterized using realistic contact mixing matrices in a social network [53] as well as with an age structure (more generally, multilayer networks [54]) can be used to optimize social network-based distancing protocol (targeted social distancing) [34]. Further work along this direction is promising and will help provide practical guidance. Moreover, it appears that instead of the actual likelihood to get infected, one’s perceived likelihood to get infected influences the decision whether to engage in social distancing and face covering [46]. Variation in individual risk assessment might therefore influence the results in our model and be an interesting extension to the model in future work.
In sum, we analyse and characterize oscillatory dynamics in the dilemma of social distancing, which arises from the non-trivial feedback between disease prevalence and behavioural intervention. Our results suggest an oscillatory tragedy of the commons in disease control when individuals act in their own right without coordination or in the absence of centralized institutions to enforce their compliance, a phenomenon that has been observed in past pandemics like the Spanish flu [55] and seems to repeat in the current COVID-19 pandemic [56]. Our work provides new insight into the dual role of human behaviour that can fuel, or fight against, the pandemic [57]. To resolve the dilemma of disease control from global pandemics to resurgence of common diseases (like measles which has become endemic in some regions [58]), a deep understanding of pertinent behavioural aspect in disease control and prevention, and large-scale human cooperation in particular, is urgently needed and will help to better inform pandemic support in the future [49].
Acknowledgements
We thank Dan Rockmore and Nicholas Christakis for helpful discussions.
Ethics
Ethical assessment is not required prior to conducting the research reported in this paper, as the present study does not have experiments on human subjects and animals, and does not contain any sensitive and private information.
Data accessibility
All the data and analysis pertaining to this work has been included in the main text. This article has no additional data.
Authors' contributions
A.G. and F.F. conceived the model; A.G. performed simulations and analyses, plotted the figures and wrote the manuscript; F.F. contributed to analyses and writing. All authors give final approval of publication.
Competing interests
We declare we have no competing interests.
Funding
F.F. is grateful for financial support by the Bill & Melinda Gates Foundation(award no. OPP1217336), the NIH COBRE Program (grant no. 1P20GM130454), the Neukom CompX Faculty Grant, the Dartmouth Faculty Startup Fund, and the Walter & Constance Burke Research Initiation Award.
Reference
- 1.Petersen LR, Jamieson DJ, Powers AM, Honein MA. 2016. Zika virus. N. Engl. J. Med. 374, 1552–1563. ( 10.1056/NEJMra1602113) [DOI] [PubMed] [Google Scholar]
- 2.Leroy EM, Kumulungui B, Pourrut X, Rouquet P, Hassanin A, Yaba P, Délicat A, Paweska JT, Gonzalez J, Swanepoel R. 2005. Fruit bats as reservoirs of Ebola virus. Nature 438, 575–576. ( 10.1038/438575a) [DOI] [PubMed] [Google Scholar]
- 3.Andersen KG, Rambaut A, Lipkin W, Holmes EC, Garry RF. 2020. The proximal origin of SARS-CoV-2. Nat. Med. 26, 450–452. ( 10.1038/s41591-020-0820-9) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 4.Lloyd-Smith JO. 2017. Infectious diseases: predictions of virus spillover across species. Nature 546, 603–604. ( 10.1038/nature23088) [DOI] [PubMed] [Google Scholar]
- 5.Bauch CT, Galvani AP. 2013. Social factors in epidemiology. Science 342, 47–49. ( 10.1126/science.1244492) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 6.Verelst F, Willem L, Beutels P. 2016. Behavioural change models for infectious disease transmission: a systematic review (2010–2015). J. R. Soc. Interface 13, 20160820 ( 10.1098/rsif.2016.0820) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 7.Fu F, Rosenbloom DI, Wang L, Nowak MA. 2011. Imitation dynamics of vaccination behaviour on social networks. Proc. R. Soc. B 278, 42–49. ( 10.1098/rspb.2010.1107) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 8.Reluga TC, Bauch CT, Galvani AP. 2006. Evolving public perceptions and stability in vaccine uptake. Math. Biosci. 204, 185–198. ( 10.1016/j.mbs.2006.08.015) [DOI] [PubMed] [Google Scholar]
- 9.Wang Z, Bauch CT, Bhattacharyya Sa, Manfredi P, Perc M, Perra N, Salathé M, Zhao D. 2016. Statistical physics of vaccination. Phys. Rep. 664, 1–113. ( 10.1016/j.physrep.2016.10.006) [DOI] [Google Scholar]
- 10.Bauch CT, Bhattacharyya S. 2012. Evolutionary game theory and social learning can determine how vaccine scares unfold. PLoS Comput. Biol. 8, e1002452 ( 10.1371/journal.pcbi.1002452) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 11.Wang W, Liu Q, Cai S, Tang M, Braunstein LA, Stanley HE. 2016. Suppressing disease spreading by using information diffusion on multiplex networks. Sci. Rep. 6, 29259. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 12.Salathé M, Bonhoeffer S. 2008. The effect of opinion clustering on disease outbreaks. J. R. Soc. Interface 29, 1505–1508. ( 10.1098/rsif.2008.0271) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 13.Funk S, Salathé M, Jansen VAA. 2010. Modelling the influence of human behaviour on the spread of infectious diseases: a review. J. R. Soc. Interface 7, 1247–1256. ( 10.1098/rsif.2010.0142) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 14.Fenichel EP. et al. 2011. Adaptive human behavior in epidemiological models. Proc. Natl Acad. Sci. USA 108, 6306–6311. ( 10.1073/pnas.1011250108) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 15.Fu F, Christakis NA, Fowler JH. 2017. Dueling biological and social contagions. Sci. Rep. 7, 43634 ( 10.1038/srep43634) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 16.Chen X, Fu F. 2019. Imperfect vaccine and hysteresis. Proc. R. Soc. B 286, 20182406 ( 10.1098/rspb.2018.2406) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 17.Bauch CT. 2005. Imitation dynamics predict vaccinating behaviour. Proc. R. Soc. B 272, 1669–1675. ( 10.1098/rspb.2005.3153) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 18.Arefin MR, Masaki T, Tanimoto J. 2020. Vaccinating behaviour guided by imitation and aspiration. Proc. R. Soc. A 476, 20200327 ( 10.1098/rspa.2020.0327) [DOI] [Google Scholar]
- 19.Cressman R, Tao Y. 2014. The replicator equation and other game dynamics. Proc. Natl Acad. Sci. USA 111, 10810–10817. ( 10.1073/pnas.1400823111) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 20.Sigmund K, Hauert C, Nowak MA. 2001. Reward and punishment. Proc. Natl Acad. Sci. USA 98, 10 757–10 762. ( 10.1073/pnas.161155698) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 21.Perc M, Jordan JJ, Rand DG, Wang Z, Boccaletti S, Szolnoki A. 2017. Statistical physics of human cooperation. Phys. Rep. 687, 1–51. ( 10.1016/j.physrep.2017.05.004) [DOI] [Google Scholar]
- 22.Page KM, Nowak MA, Sigmund K. 2000. The spatial ultimatum game. Proc. R. Soc. Lond. B 267, 2177–2182. ( 10.1098/rspb.2000.1266) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 23.Capraro V, Perc M, Vilone D. 2020. The evolution of lying in well-mixed populations. J. R. Soc. Interface 16, 20190211 ( 10.1098/rsif.2019.0211) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 24.Kumar A, Capraro V, Perc M. 2020. The evolution of trust and trustworthiness. J. R. Soc. Interface 17, 20200491 ( 10.1098/rsif.2020.0491) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 25.Capraro V, Perc M. 2019. Grand challenges in social physics: in pursuit of moral behavior. Front. Phys. 6, 107 ( 10.3389/fphy.2018.00107) [DOI] [Google Scholar]
- 26.Chen X, Fu F. 2018. Social learning of prescribing behavior can promote population optimum of antibiotic use. Front. Phys. 6, 139 ( 10.3389/fphy.2018.00139) [DOI] [Google Scholar]
- 27.Townsend AK, Hawley DM, Stephenson JF, Williams KE. 2020. Emerging infectious disease and the challenges of social distancing in human and non-human animals. Proc. R. Soc. B 287, 20201039 ( 10.1098/rspb.2020.1039) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 28.Nicola M, Alsafi Z, Sohrabi C, Kerwan A, Al-Jabir A, Iosifidis C, Agha M, Agha R. 2020. The socio-economic implications of the coronavirus and COVID-19 pandemic: a review. Int. J. Surg. 78, 185–193. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 29.Reluga TC. 2010. Game theory of social distancing in response to an epidemic. PLoS Comput. Biol. 6, e1000793 ( 10.1371/journal.pcbi.1000793) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 30.Maharaj S, Kleczkowski A. 2012. Controlling epidemic spread by social distancing: do it well or not at all. BMC Public Health 12, 679 ( 10.1186/1471-2458-12-679) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 31.Huberts N, Thijssen J. 2020. Optimal Timing of Interventions during an Epidemic. Available at SSRN 3607048.
- 32.Caley P, Philp DJ, McCracken K. 2008. Quantifying social distancing arising from pandemic influenza. J. R. Soc. Interface 5, 631–639. ( 10.1098/rsif.2007.1197) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 33.Wallinga J, van Boven M, Lipsitch M. 2010. Optimizing infectious disease interventions during an emerging epidemic. Proc. Natl Acad. Sci. USA 107, 23–928. ( 10.1073/pnas.0908491107) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 34.Aleta A. et al. 2020. Modelling the impact of testing, contact tracing and household quarantine on second waves of COVID-19. Nat. Hum. Behav. 4, 964–971. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 35.Kissler SM, Tedijanto C, Goldstein E, Grad YH, Lipsitch M. 2020. Projecting the transmission dynamics of SARS-CoV-2 through the postpandemic period. Science 368, 860–868. ( 10.1126/science.abb5793) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 36.Sethi SP, Staats PW. 1978. Optimal control of some simple deterministic epidemic models. J. Oper. Res. Soc. 29, 129–136. ( 10.1057/jors.1978.27) [DOI] [Google Scholar]
- 37.Abakuks A. 1974. Optimal immunisation policies for epidemics. Adv. Appl. Probab. 6, 494–511. ( 10.2307/1426230) [DOI] [Google Scholar]
- 38.Abakuks A. 1973. An optimal isolation policy for an epidemic. J. Appl. Probab. 10, 247–262. ( 10.2307/3212343) [DOI] [Google Scholar]
- 39.Wickwire KH. 1975. Optimal isolation policies for deterministic and stochastic epidemics. Math. Biosci. 26, 325–346. ( 10.1016/0025-5564(75)90020-6) [DOI] [Google Scholar]
- 40.Morris DH, Rossine FW, Plotkin JB, Levin SA. 2020 Optimal, near-optimal, and robust epidemic control. (http://arxiv.org/abs/2004.02209. )
- 41.Simon HA. 1990. Bounded rationality. In Utility and probability pp. 15–18. New York, NY: Springer.
- 42.Tversky A, Kahneman D. 1992. Advances in prospect theory: cumulative representation of uncertainty. J. Risk Uncertain. 5, 297–323. ( 10.1007/BF00122574) [DOI] [Google Scholar]
- 43.Weitz JS, Eksin C, Paarporn K, Brown SP, Ratcliff WC. 2016. An oscillating tragedy of the commons in replicator dynamics with game-environment feedback. Proc. Natl Acad. Sci. USA 113, E7518–E7525. ( 10.1073/pnas.1604096113) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 44.Tilman AR, Plotkin JB, Akçay E. 2020. Evolutionary games with environmental feedbacks. Nat. Commun. 11, 1–11. ( 10.1038/s41467-020-14531-6) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 45.Wang X, Zheng Z, Fu F. 2020. Steering eco-evolutionary game dynamics with manifold control. Proc. R. Soc. A 476, 20190643 ( 10.1098/rspa.2019.0643) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 46.Capraro V, Barcelono H. 2020. The effect of messaging and gender on intentions to wear a face covering to slow down COVID-19 transmission. J. Behav. Econ. Policy 4, 45–55. [Google Scholar]
- 47.Jordan JJ, Yoeli E, Rand DG. 2020 Don’t get it or don’t spread it? Comparing self-interested versus prosocial motivations for COVID-19 prevention behaviors. (http://arxiv.org/abs/10.31234/osf.io/yuq7x. )
- 48.Van Bavel J.et al.2020National identity predicts public health support during a global pandemic. (http://arxiv.org/abs/10.31234/osf.io/ydt95)
- 49.Van Bavel JJ. et al. 2020. Using social and behavioural science to support COVID-19 pandemic response. Nat. Hum. Behav. 4, 460–471. [DOI] [PubMed] [Google Scholar]
- 50.Apolloni A, Poletto C, Colizza V. 2013. Age-specific contacts and travel patterns in the spatial spread of 2009 H1N1 influenza pandemic. BMC Infect. Dis. 13, 176 ( 10.1186/1471-2334-13-176) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 51.Eames, KDT, Tilston NL, Brooks-Pollock E, Edmunds WJ. 2012. Measured dynamic social contact patterns explain the spread of H1N1v influenza. PLoS Comput. Biol. 8, e1002425 ( 10.1371/journal.pcbi.1002425) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 52.Wu JT, Leung K, Bushman M, Kishore N, Niehus R, de Salazar PM, Cowling BJ, Lipsitch ML, Leung GM. 2020. Estimating clinical severity of COVID-19 from the transmission dynamics in Wuhan, China. Nat. Med. 26, 506–510. ( 10.1038/s41591-020-0822-7) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 53.Block P, Hoffman M, Raabe IJ, Dowd JB, Rahal C, Kashyap R, Mills MC. 2020. Social network-based distancing strategies to flatten the COVID-19 curve in a post-lockdown world. Nat. Hum. Behav. 4, 583–596. [DOI] [PubMed] [Google Scholar]
- 54.Aleta A, de Arruda GF, Moreno Y. 2020 Data-driven contact structures: from homogeneous mixing to multilayer networks. (http://arxiv.org/abs/2003.06946. )
- 55.Taubenberger JK, Morens DM. 2006. 1918 Influenza: the mother of all pandemics. Emerg. Infect. Dis. 12, 15–22. ( 10.3201/eid1209.05-0979) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 56.Alwan NA. et al. 2020. Scientific consensus on the COVID-19 pandemic: we need to act now. Lancet 396, e71–e72. ( 10.1016/S0140-6736(20)32153-X) [DOI] [PMC free article] [PubMed] [Google Scholar]
- 57.2020 Behaviour fuels, and fights, pandemics. See https://www.nature.com/articles/s41562-020-0892-z (accessed on 24 June 2020).
- 58.Muscat M. et al. 2009. Measles in Europe: an epidemiological assessment. Lancet 373, 383–389. ( 10.1016/S0140-6736(08)61849-8) [DOI] [PubMed] [Google Scholar]
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