The objective of this study is to gain insight into the role of human behavior in 69 modulating the spread and prevalence of COVID-19. We construct a mathematical 92 We construct a model of COVID-19 transmission with quarantine and hospitalization 93 in Section 4.1. We follow the natural history of the infection [42, 53] and partition 94 the population according to their disease status as susceptible (S(t)), exposed (E(t)), 95 asymptomatically infected (A(t)), symptomatically infected (I(t)), quarantined (Q(t)), 96 hospitalized (H(t)), and removed (R(t)) individuals. The flow diagram depicting the 97 transition from one state to the other as the disease progresses through the population 98 is shown in Figure 1 , and the associated state variables and parameters are described 99 in Tables 1. 3 . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
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Proportion of susceptible individuals E(t) Proportion of exposed individuals A(t)
Proportion of asymptomatically infected individuals I(t)
Proportion of symptomatically infected individuals Q(t)
Proportion of hospitalized individuals R(t)
Proportion of removed individuals Parameter Description β Infection rate η A , η Q , η H Modification parameters for asymptomatic, quarantined, and hospitalized infection rates q Proportion of exposed developing asymptomatic infections σ Disease progression rate from the exposed to infectious γ I , γ A , γ Q , γ H Recovery rates of symptomatic, asymptomatic, quarantined, and hospitalized individuals ω Q , ω H Quarantine and hospitalization rates ν Q Quarantine violation rate ν H Hospital discharge rate δ I , δ A , δ Q , δ H Death rates of symptomatic, asymptomatic, quarantined, and hospitalized individuals
The associated reproduction number [22, 48] of the baseline COVID-19 model (1) 101 with quarantine and hospitalization, denoted by R 0 , is given by
where 103 R I = (1 − q)β(k 3 η H ω H + k 4 η Q ω Q + k 3 k 4 ) (k 2 k 3 k 4 − k 3 ν H ω H − k 4 ν Q ω Q ) and R A = qβη A k 1 , with k 1 = γ A + δ A , k 2 = γ I + ω Q + ω H + δ I , k 3 = ν Q + γ Q + δ Q , and k 4 = ν H + γ H + δ H .
The quantity R I is the number of secondary infections produced by symptomatic in-105 dividuals, while R A is the number of secondary infections generated by asymptomatic 106 individuals. Together, the epidemiological quantity R 0 , measures the average number 107 of COVID-19 secondary infections produced when a single infected individual is in-108 troduced into a completely susceptible population [22, 48] . Hence, COVID-19 can be 109 effectively controlled in the population if the reproduction number (R 0 ) can be reduced 110 to (and maintained at) a value less than unity (i.e., R 0 < 1). 111 We computed the numerical value of the reproduction number R 0 using the pa-112 rameter values tabulated in Table 3 . Some of the parameter values in Table 3 were 113 fitted based on the COVID-19 data for Arizona from January 26 to July 6 [31] (see 114 Figure 12 ), while others were obtained from the literature. Using these parameter 115 estimates, we obtain R 0 ≈ 1.84 for the COVID-19 outbreak in Arizona. does not depend on the quarantine and hospitalization rates (see also Figure 13 (a)).
Using this lower level of the infection rate, we see in Figure 13 (b) that R 0 can be kept 128 below 1 provided either the quarantine rate is above 0.4 or the hospitalization rate is 129 above 0.2. Varying quarantine rate ω Q and hospitalization rate ω H . (b) Varying quarantine rate ω Q and infection rate β.
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint the hospitals into the community due to overwhelmed demand for hospitalizations or 132 lack of resources, then the disease burden is high and containing the disease becomes 133 challenging as values of R 0 are greater than 1 for all values of ν Q and ν H (see Fig-134 ure 3(a)). The situation is even worse if quarantine violation is varied along with 135 poor hygiene and disregard for social distancing, which increases the infection rate β. Table 2 . . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity. (which was not certified by peer review)
The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint socio-economic losses due to the partial economy shutdown; the socio-economic losses 182 accumulate over time. We assumed that the decision to enact appropriate closures 183 stays in effect if and only if a certain minimum time has passed since the start of the 184 pandemic and at least half of the susceptible population supports closures.
In all simulations involving susceptible individual support for closure, we assume 186 that the effectiveness of closures is C 0 = 0.6 and the initial time the closure decision 187 may be enacted is t close = 30 days. Figure 5 the closure support drops below the threshold, which results in (partial) re-openings.
However, the number of infected individuals is still relatively high, and a second bigger 208 8 . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity. (which was not certified by peer review)
The copyright holder for this preprint this version posted September 2, 2020. to protect others. The initial predisposition of the population to the altruistic act of 236 self-isolation determines the peak of the epidemic and its timing ( Figure 6 (a)). The 237 more individuals are willing to self-isolate, the lower the peak and the later it occurs. 238 We considered one set of fixed values of the symptomatically infected individual 239 social learning rate parameter κ I and the sensitivity to self-isolation parameter ε I . 240 We investigate the effects of varying these parameters in a full behavioral model. In . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. In general, we assumed that κ S < κ I because supporting school and workplace 276 10 . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. that is why we assumed that the social learning rate κ S for closure support behavior 282 is lower than that for self-isolating behavior (κ I ).
283 Figure 8 shows that increasing the support closure behavior social learning rate . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint to the epidemiological situation, individuals switch back to non-compliance as soon as 299 the situation improves but well before the disease prevalence is reduced to negligible 300 numbers. This, in turn, results in a new spike of infections. We note that this phe- causes multiple oscillations in self-isolating behavior (Figure 11(b) ) and hence future 320 waves of infection.
The take home-message from the results presented in Figures 10 and 11 is that 
(iv) It is important to incentivize the cost and burden of self-isolation to encour-337 age more symptomatic individuals to self-isolate because high sensitivity to self-338 isolation is not beneficial to the community as a whole. CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint turned out that the peak of the epidemic curve generated by symptomatic infections 374 willing to self-isolate was lower than the peak of the epidemic curve generated by the 
Our key findings further show that when the symptomatic infectious population 392 learn the positive behavior or are more willing to self-isolate, the community benefits, 393 even though this change in behavior comes at a cost to them. Self-isolation often comes 394 with financial implications and distress; not very many people can bear these burdens.
Hence, it is important to incentivize self-isolation of the symptomatic infectious pop-396 ulation as many infected people will rather stay home than go to work since staying 397 at home will help the public good and create an opportunity to help save more lives 398 [26] . One way to incentivize the symptomatic infectious is to pay them to stay home, 399 perhaps via direct government subsidies for sick leave for infected individuals [26] . Our CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
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The goal of this study was to provide insight into possible effects of human behavior on 421 non-pharmaceutical intervention strategies (such as partial lock-down and social isola- CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
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In this section, we use the imitation dynamic approach of evolutionary game theory 
Following [41], we define 523 C(t) = 0 if t < t close or x S < 1/2 C 0 if t ≥ t close and x S ≥ 1/2
(3)
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint where C 0 is a combined measure of the effectiveness of physical distancing in those 524 workplaces that remain open and how many workplaces are closed. The decision to 525 close schools and workplaces is "turned on" if the time after the start of the pandemic 526 is at least t close and at least half of the (susceptible) population supports closure. The 527 closure policy is "turned off" if less than half of the (susceptible) population supports 528 closure.
The susceptible individuals weigh the risk of the infection based on the disease 530 prevalence and the accumulating socio-economic losses due to the closures. The sus-531 ceptible individuals who do not support school and workplace closure are willing to 532 face the risk of infection, and their perceived payoff is given by
where π S is the sensitivity to being infected with COVID-19 parameter. The susceptible 534 individuals who support closure efforts face socio-economic losses, and their perceived 535 payoff is given by
where ρ S is the the sensitivity to the accumulated socio-economic losses L S (t), as in 537
[41]. 538 We now describe how the behavioral responses of susceptible individuals evolve with 539
time. An individual who did not support closure but decided to switch its strategy 540 achieves a payoff gain
We assume that individuals employ a social learning process where they adopt strate-542 gies of other individuals with the rate proportional to the payoff gain, which can be 543 realized via an imitation dynamic. The proportion of susceptible individuals who sup-544 port closure thus evolves according to
where κ S is the social learning rate. The individuals who do not support closure (1−x S ) 546 sample the individuals who do support closure (x S ) and switch their strategy at the 547 rate proportional to the payoff gain ∆E S . Using equation (6), we obtain Since scaling payoff functions does not affect the outcome, we can replace ∆E S 553 given by (6) with ∆E S = I(t) + Q(t) + H(t) − (ρ S /π S )L S (t). Then
20 . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint where ε S = ρ S /π S is the sensitivity to the socio-economic losses relative to getting 555 infected with COVID-19. 556 Finally, following [41] , the evolution of the time-varying quantity L S (t), which 557 represents the accumulated socio-economic losses, obeys the exponential fading memory 558 mechanism given by
where α S controls the rate at which school and workplace closures impacts socio-560 economic health of the population, and ξ S is a decay rate that represents adjustment 561 to the baseline losses. 
We assume that the symptomatically infected individuals who tested positive for 577 COVID-19 and were ordered to quarantine themselves leave quarantine at a constant 578 rate ν Q . However, symptomatically infected individuals (I(t)) whose condition was not 579 severe enough to go to a hospital and/or get tested may elect to self-isolate to protect 580 others. Let x I (t) be the proportion of symptomatically infected individuals I(t) who 581 elect to self-isolate. We assume that self-isolated individuals do not contribute to 582 the spread of the infection, and the force of infection term involving I(t) becomes
(1 − x I (t))I(t). Hence, the equations for the susceptible and exposed individuals from 584 the baseline model become
A symptomatically infected individual who elects not to self-isolate faces the burden 586 of infecting other individuals. These individuals use the publicly available information 587 on the COVID-19-induced death rates to estimate the extent of the burden. We 588 therefore assume that the payoff of an individual who chooses not to self-isolate is 589 given by
where π I is the sensitivity to infecting others parameter. On the other hand, an infected 591 individual who decides to self-isolate faces a fixed cost of such a decision because the 592 21 . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity.
The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint length of self-isolation is approximately equal to the time it takes to recover. Hence, 593 the payoff of an individual who chooses to self-isolate is given by
where ρ I is the sensitivity to self-isolation parameter.
Similar to the closure support model described above, the proportion of symptomat-596 ically infected individuals who elect to self-isolate evolves according to the imitation
where κ I is the self-isolation social learning rate, and ε I = ρ I /π I is the sensitivity 599 to self-isolation relative to infecting others. The (conscientious) infected individuals 600 would tend to self-isolate if the COVID-19-induced death toll is high, while they would 601 tend not to self-isolate as long as the death rates become sufficiently low. system is given by the following system of ordinary differential equations:
The game-theoretic model of dynamic human behavior state variables and parameters 612 are summarized in Table 2 .
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint quarantine and buy alcohol. 688 https: // www. theguardian. com/ world/ 2020/ jul/ 10/ . CC-BY-NC 4.0 International license It is made available under a is the author/funder, who has granted medRxiv a license to display the preprint in perpetuity. (which was not certified by peer review)
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint (ε I ) on symptomatic infectious 840 Here we show the simulation results for all symptomatic infections with high sensitivity 841 to self-isolation ε I = 0.00008. 
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The copyright holder for this preprint this version posted September 2, 2020. . https://doi.org/10.1101/2020.08.30.20184804 doi: medRxiv preprint cial learning rates κ I
Here we show the simulation results of varying symptomatically infected individuals 845 social learning rate κ I with high sensitivity to self-isolation ε I = 0.00008. 
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