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c Founder, covid-testing.org
Toronto, Canada
Abstract
around the world in the fight against COVID-19: Social distancing, shelter-in-place,
mask wearing, etc measures to protect the susceptible, together with, in various degrees,
testing & contact-tracing to identify, isolate and treat the infected. The majority of
countries have relied on the former, while ramping up their testing and tracing
capabilities. We consider the examples of South Korea, Italy, Canada and the United
States. By fitting a disease transmission model to daily case report data, we show that
to date have had a significant impact on the evolution of their pandemic curves. In this
work we estimate the average isolation rates of infected individuals needing to occur in
each country as a result of large-scale testing and contact tracing as a mean of lifting
isolation rate of an infected individual every 4.5 days (South Korea), 5.7 days (Canada)
and to 6 days (Italy) would be sufficient. We also find that a rate of under 3.5 days will
help in the United States, although it would not completely mitigate the second wave
¶Edward W. Thommes, Laurent Coudeville and Ayman Chit are employees of Sanofi
Pasteur. Monica G. Cojocaru has received national research grants in the past in which
Sanofi Pasteur was a matching partner. The grants are for research completely
1 Introduction 1
In late 2019, a novel betacoronavirus called SARS-CoV-2 emerged from a live animal 2
marketplace in Wuhan, Hubei Province, China, and has since inflicted a worldwide 3
with an estimated R0 between 2.2 and 4.6 [1, 2] although it is important to consider that 5
behavioural and environmental factors [3]. The incubation period has been found to be 7
5.1-5.2 days, while 97.5% of patients display symptoms within 11.5 days [1, 4]. The 8
disease spreads primarily through the respiratory tract and respiratory secretions. 9
As of July 16 2020, there have been a total of 13.7 million confirmed cases of 10
COVID-19 worldwide, and over 580,000 deaths (WHO COVID-19 Situation Report 89, 11
https://www.who.int/emergencies/diseases/novel-coronavirus-2019/ 12
active, with new cases reported daily, though a number of countries have clearly passed 14
a (first) peak. Countries have taken various degrees of social distancing measures: 15
wearing etc. (e.g. [5–8]) in an effort to suppress the disease, or at least prevent it from 17
overwhelming a country’s critical care capacity. While proven effective to slow the 18
spread, these measures have had a large effect on daily lives and economies throughout 19
the world. Countries who managed to stave off their first wave, are now in the process 20
further examine and devise strategies which will allow more phasing out of social 23
In this work, we fit a disease transmission model to daily case reports in four 25
We begin with the examples of Italy, Canada and the United States, three countries 29
which have relied principally on social distancing via universal shelter-in-place measures, 30
while testing and contact tracing was implemented at an increased pace only after the 31
evolution scenarios to that of South Korea, a country which has had and still has a tight 33
can still provide, from this moment onward, a feasible path to further relaxing 36
pandemic profiles. The case of the United States stands apart: while coordinated large 38
scale, frequent testing and contact tracing will help decelerate the current U.S. 39
The structure of the paper is as follows: in Section 2 we introduce our model main 42
ideas, notation, and assumptions. We follow in Section 3 with the presentation of our 43
fitted infection curves for Italy, Canada, the U.S., and South Korea, wherein we infer 44
the net effect to date of social distancing, testing & contact tracing on the decrease of 45
the transmission rates in each of the 4 countries. We then evaluate scenarios for 46
outbreaks by assuming an increase in testing and contact tracing. We are able to derive 48
the frequency with which a tested (infected) individual and an exposed (traced) 49
individual need to be detected and isolated in order for each country to maintain an 50
effective reproduction number of 1 (that is to say, each country maintains a ”slow burn” 51
of the form: 61
ds
= −βsi (1)
dt
de
= βsi − σe
dt
di
= σe − γi
dt
dr
= γi
dt
S E I R
with s = N, e= N, i= N, r= N, s + e + i + r = 1 and where β is the rate of effective
contacts, 1/σ = Tlat is the mean latent period (which may differ from the incubation
period), and 1/γ = Tinf is the mean duration of infectiousness, with both times having
exponential distributions. We also have the auxiliary equation for the cumulative
number of cases,
dc
= σe
dt
inci = ci − ci−1
In turn, a SIR model is similar to (1) but without the E compartment, thus only 3 63
The spread of an infectious disease can be halted if its effective reproduction number 65
Ref f = R0 s, (2)
can be decreased below 1. The effective reproduction number of both the SIR and SEIR 66
β
Ref f = s. (3)
γ
In both the SIR and SEIR model, when s ≈ 1 and we are near the disease-free 68
equilibrium (1, 0, 0, 0), the early growth of both i and inc is exponential (see e.g. [12]): 69
ρSIR + γ
R0SIR = (6)
γ
p
−(σ + γ) + (σ − γ)2 + 4σβ
ρSEIR = (7)
2
(ρSEIR + σ)(ρSEIR + γ)
R0SEIR = . (8)
σγ
In the limit as σ → ∞, the SEIR model reduces to the SIR model, and accordingly, as 74
For COVID-19, as for other pandemics (e.g. SARS, MERS, the 1918 Spanish flu), 76
we can assume the entire population to be initially susceptible. Therefore, in the early 77
stages of an outbreak, Ref f ≈ R0 . We will also assume that infection with COVID-19 78
confers subsequent immunity, which does not wane significantly over the time horizon 79
considered here. Thus, whether they die or recover, an infected person is considered 80
removed from the pool of susceptibles. In the absence of a vaccine or other control 81
measures, s(t) = 1 − c(t), where c(t) is the cumulative number of people infected at 82
time t. 83
From Equation 3, assuming β to be given, we see that Ref f can be decreased in two 84
ways: by decreasing s at a rate higher than that due to infection alone; or by increasing 85
these effectively take a part of the population “out of circulation” as far as disease 87
To explicitly depict the role of control measures, we adapt the SEIR model to a 90
auxiliary equation for C, the cumulative number of infected. The resulting SEIRL 92
ds
= −βsi
dt
de
= βsi − σe − κ1 e
dt
di
= σe − (γ + κ)i
dt
dr
= γi
dt
dl
= κi + κ1 e
dt
dc
= σe
dt
where as in the standard SEIR model, β is the mean rate of effective contacts, 94
1/σ = Tlat is the mean latent period, and 1/γ = Tinf is the mean infectious period. 95
Finally, 1/κ1 = Tisol,lat and 1/κ = Tisol,inf are the mean times for the latent and 96
tracing. 98
βσ
Ref f,SEIRL = s (10)
(σ + κ1 )(γ + κ)
It can be shown (see Appendix A) that the exponential growth rate (Equation 4) of 100
p
−(σ + κ1 + γ + κ) ((σ + κ1 ) − (γ + κ))2 + 4σβs
ρSEIRL = + ,
2 2
We can also express the effective reproduction number in terms of ρSEIRL : 103
3 Results 104
log(inc(t)) ∝ ρt (11)
i.e. a log-linear plot of incidence versus time will have slope ρ. Indeed, early exponential 107
growth can be seen to be a near-universal feature in COVID-19 daily case count data 108
from around the world. Figures 1 and 2 plot log(inc) versus time for South Korea, Italy, 109
Canada and the U.S., using time series data of daily new cases compiled by the Johns 110
Hopkins University Center for Systems Science and Engineering (JHU CSSE) [13], 111
http://github.com/CSSEGISandData/COVID-19/tree/master/csse_covid_19_data 113
In all four countries the initial linear phase is clearly apparent, followed by a 114
transition to sub-exponential growth. This transition is sharpest for South Korea, where 115
growth switches abruptly to decay around 1 March. Regression fit results for ρ and the 116
ln(2)
Tdbl = (12)
ρ
Fig 1. Log-linear plots of daily COVID-19 incidence versus time for South Korea and
Italy. The initial linear phase corresponds to exponential growth, which subsequently
turns over into sub-exponential growth. The factor ρ and the corresponding doubling
time are estimated via a regression fit to the initial phase. R0 is calculated using Eq. 8
with σ = γ = (2.5d)−1 .
together with dates for the onset of major national-level protective measures, are given 118
in Figures 1 and 2 and Table 1. In all four cases, the transition to sub-exponential 119
growth occurred at or after the time that widespread protective measures were first 120
invoked. 121
Inferring R0 from ρ requires choosing values for the mean latent and infectious 122
Estimates of the serial interval of COVID-19 range from 3.95 to 6.6 days [14–17]. We 124
adopt a value of Tser = 5 days. The latent period of the disease is not well constrained, 125
but it can be shown (Appendix A) that for a given value of Tser and ρ, the maximum 126
value of R0 is obtained when Tlat = Tinf = Tser /2. We assume this “worst-case” 127
We begin with the remark that all 4 countries have enacted social distancing via school 131
closures, nationwide shutdowns, shelter-in-place orders, mask wearing, etc., all in 132
Table 1. Initial exponential growth rates and R0 values for South Korea,
Italy, Canada, and the U.S.
Country ρ initial growth R0
S. Korea 0.22(0.15,0.29) 2.38(1.88,2.94)
Italy 0.18(0.16,0.2) 2.1(1.98,2.22)
Canada 0.18(0.16,0.2) 2.11(1.97,2.26)
U.S. 0.3(0.27,0.33) 3.07(2.81,3.35)
counter-measures sub-exponential onset fraction of cases reported
S. Korea Feb 21 [18] March 1 0.84
Italy March 12 [19] March 22 0.11
Canada March 20 [20] April 3 0.26
U.S. March 13 [21] April 6 0.51
Estimates of initial exponential growth rate ρ are obtained from regression fits to the
early outbreak phases (Figs 1 and 2). Corresponding values of R0 assume
Tlat = Tinf = 2.5 days. Estimates of the fraction of cases reported are taken from [22].
various combinations. South Korea was the first to impose measures, followed by Italy, 133
where the measures were ordered and coordinated eventually country-wide. In Canada 134
most provinces enacted similar measures over the course of 1-2 weeks around March 20, 135
2020, while the United States has had the more heterogeneous spread of similar 136
We interpret the transition to sub-exponential growth (Figs 1 and 2) as the first 138
signature of the effect of these measures in a given country, and use this as the starting 139
point to infer the net effect that these measures have had up to the end of June 2020. 140
For each country, we fit the SEIR model solutions for daily incidence 141
{incmodel,1 , ..., incmodel,n } to daily case reports. The model output is multiplied by a 142
obtained using delay-adjusted case fatality rates [22], and f is the fraction of cases 144
which are symptomatic, estimated to be f = 0.5, from a recent CDC report 1 . 145
We compute − log L, the normal negative log likelihood of the time series of 146
observed daily incidences, {incobs,1 , ..., incobs,n }, given the model output, as a function 147
x = (i0 , q1 , q2 , ..., qm )
where i0 is the initial number of infected and the qi are reduction factors on the rate of 149
disease transmission, varying over time, such that βi0 = qi β (see Table 1). R0 for each 150
country is fixed at the respective values obtained via regression above. Parameters are 151
drawn using unweighted (uniform) Latin hypercube sampling. The best-fit solution is 152
We present our best fits for Italy, Canada, the U.S. and South Korea in the next 154
• Linear and semi-log plots of daily incidence data of confirmed cases per country, 156
• Inferred true number of infected, taking into account under-reporting and 158
asymptomatic cases (“with measures, all”). Shown for comparison are the number 159
of confirmed cases (“no measures, confirmed”) and all cases (“no measures, all”) 160
planning-scenarios-h.pdf
• Cumulative incidence and inferred reduction in effective contact rates (i β) due to 162
• These fits assume κ = κ1 = 0, in other words, the reduction in the effective 164
contact rates qi0 s are a measure of each country’s combination of measures to date, 165
Fig 3. (A) Left two panels: Linear and semi-log plots of daily incidence data of
confirmed cases in Italy, together with maximum-likelihood model fit (“with measures,
confirmed”) Also shown is the inferred true number of infected, taking into account
under-reporting and asymptomatic cases (“with measures, all”). Shown for comparison
are the number of confirmed cases (“no measures, confirmed”) and all cases (“no
measures, all”) expected to have occurred in the absence of countermeasures. (B)
Right two panels: Cumulative incidence (top) and inferred reduction of effective
contacts, together with the corresponding effective reproduction numbers (vertical
numbers) (bottom).
In all four countries, interventions arrested the initial exponential rise in cases and 167
brought the effective reproduction number below. In Italy, South Korea and Canada, 168
daily case numbers have since been brought far below their peak values. 169
South Korea effected the strongest and most rapid reduction in transmission. South 170
Korea experienced a very similar early exponential growth in cases, and hence has a 171
similar inferred R0 , as the other three countries. However, its mitigation and control 172
measures stood out from the beginning in the fact that the country employed a rapid 173
scale-up of testing, concurrent with contact tracing and isolating of infected individuals. 174
There are also social distancing measures imposed, but notably no shelter-in-place. Last 175
but not least, mask wearing is a regular policy that the population adopts widely (bot 176
Fig 5. U.S. daily COVID-19 data and model fit for incidence and cumulative incidence;
see caption of Fig 3 for details
only for this pandemic, but also for flu for example). Members of the population also 177
contrast, mask wearing was instituted much later in the other 3 countries and some 179
After some relaxation of measures in South Korea, together with a series of national 181
holidays from April 30 to May 5 (“Golden Week”) possibly playing a role, a resurgence 182
occurred. This appears to have since been stabilized, with (see Fig 1 above). Our fit is 183
Fig 6. South Korea daily COVID-19 data and model fit for incidence and cumulative
incidence; see caption of Fig 3 for details
As presented in Section 2, for a given set of values of β, σ and γ, Equation 10 gives us a 189
closed form expression for Ref f,SEIRL as a function of κ and κ1 . This relationship is 190
1
depicted as a surface plot in Fig 7 for σ = γ = 2.5 and σ = γ = 15 . In the latter case, 191
Ref f,SEIRL is nearly twice as large as in the former. It is interesting to note, though, 192
that for both cases, the combinations of κ and κ1 that make Ref f,SEIRL = 1 (i.e. the 193
intersections of the respective surfaces with the R0 = 1 plane) are quite similar. This 194
respectively, it is the latter which increasingly dominate the rate at which 196
Fig 7. Effects of isolation rates due to testing and contact tracing on the initial value
1
of R0S(Q)EIRL model. We computed σ = γ = 2.5 (upper most surface), σ = γ = 15
(middle surface) and the reference surface R0 = 1.
From Equation 10, we obtain the relationship between κ and κ1 that makes 198
sβσ sβσ
1= =⇒ k = −γ (14)
(σ + κ1 )(γ + κ) σ + κ1
Extracting the current values of s from our simulations of last section, we can now 200
compute average isolation rates due to testing and tracing that would ensure an 201
effective reproduction number of 1, while assuming that the effective contact rates β 202
will reverse to their values near the disease-free equilibrium, before any other social 203
distancing measures were employed. We further assume that the isolation rate due to 204
contact tracing ( κ1 ) is the same as the isolation rate due to testing (κ) and we used 205
Table 2. Model results for South Korea, Italy, Canada, and the U.S. for
Ref f = 1.
1 1
β s Ref f κ = κ1 ( , )
κ κ1
SK 0.961 0.999 1 0.22 (4.5, 4.5) days
Italy 0.841 0.937 1 0.161 (6.2,6.2) days
Can 0.841 0.982 1 0.175 (5.7,5.7) days
U.S. 1.225 0.96 1 0.288 (3.5, 3.5) days
1
In all cases, we chose γ = σ = 2.5 for a Tser = 5 days and where s are the estimated
values of susceptibles remaining in each country in mid July 2020.
While the U.S. and South Korea do not seem to have an effective reproduction 207
number under 1 at the moment, Canada and Italy have their effective reproductive 208
number estimated to Ref f = 0.83, as seen from Figures 1, 2. In their cases, we can redo 209
our estimates for the 2 countries and compute the isolation rates due to testing and 210
tracing so that they maintain their current value of Ref f = 0.83 (see Table 3): 211
Table 3. Model results for South Korea, Italy, Canada, and the U.S. for
Ref f = 0.83.
1 1
β s Ref f κ = κ1 ( , )
κ κ1
Italy 0.841 0.937 0.83 0.216 (4.62,4.62) days
Can 0.841 0.982 0.83 0.23 (4.33,4.33) days
1
In all cases, we chose γ = σ = 2.5 for a Tser = 5 days and where s are the estimated
values of susceptibles remaining in each country in mid July 2020. We used Eq (8).
We present next pandemic forecasts under different testing and contact tracing rates, in 213
the four countries under consideration. We show how the theoretical estimates arise in 214
the context of the simulated pandemic evolution in each of the 4 countries. 215
We depict first Canada and Italy, as they have with similar estimates, in several 216
Fig 8. Predicted daily cases in the Canada under different rates of isolation due to
testing, or testing plus contact-tracing, accompanied by a cessation of distancing
measures.
thresholds given in Table 2, a second pandemic wave is averted in each respective 219
country. This threshold is the least stringent in Italy, where about 6% of the population 220
(accounting for asymptomatic and/or unreported cases) is inferred to have been infected 221
Next we present the simulations for South Korea and the United States: We see 223
that the isolation rates are the most stringent (lowest values in days) for the U.S., where 224
the value of R0 inferred from the initial exponential rise of cases is right now higher 225
than that of the other three countries (≈ 1.2). In the case of the United States, a 226
large-scale testing and tracing operation alone will not be able to curtail the current 227
epidemic curve around, thus strong social-distancing measures will be still be needed. 228
Current testing guidelines for social-distancing relaxation measures are established 229
by the WHO in such a way that countries can relax these if positivity rates for testing 230
are under 5% for 14 days in a row. Currently, South Korea is at 1.08%, Italy = 2.39%, 231
2
Canada = 4.34% and U.S. = 6.28% From publicly available data we have that daily 232
testing (in numbers per day/per 1 million) are now as: Italy = 0.09823, 233
Canada=0.085104, U.S.=0.1282 3 . We note that all these rates are lower than what we 234
2 https://coronavirus.jhu.edu/testing/international-comparison
3 https://www.statista.com/statistics/1104645/covid19-testing-rate-select-countries-worldwide/
Fig 9. Predicted daily cases in Italy under different rates of isolation due to testing, or
testing plus contact-tracing, accompanied by a cessation of distancing measures
accompanied by different with various isolation rates due to testing only, or a
combination of testing and contact tracing
require in our tables above, however this is consistent with the fact that at the moment 235
none of South Korea, Canada, Italy or U.S. have completely removed all 236
social-distancing measures. These values give us an idea of a possible current isolation 237
1 1
rate due to testing/tracing: 0.1282 = 7.8 days for the U.S. and of 0.085 = 11 days for 238
Canada. 239
5 Discussion 240
In keeping with other published findings for these and other countries, our results 241
suggest that the COVID-19 countermeasures taken in South Korea, Italy, Canada and 242
the United States have had a substantial impact on the course of the disease. Even 243
accounting for estimated under-reporting, the number of cases in these countries 244
appears thus far to have been suppressed by roughly one order of magnitude in Italy 245
and the U.S., two orders of magnitude in Canada, and three orders of magnitude in 246
South Korea. The development of effective vaccines and treatments is still critical to the 247
future control of this disease, however in the interim, non-pharmaceutical interventions 248
Modeling studies and the United States case show that, barring a proportion of 250
Fig 10. Predicted daily cases in the South Korea under different rates of isolation due
to testing, or testing plus contact-tracing, accompanied by a cessation of distancing
measures.
asymptomatic cases so large that the majority of people have already been infected, a 251
second wave of disease is inevitable if distancing measures are halted or relaxed early. 252
As shown in the case of the US, it is vital not to rush into a relaxing of distancing 253
measures. As illustrated in Figures 9 to 10, if a change in control strategy causes Ref f 254
to exceed 1, how quickly a second wave builds depends on the number of cases at the 255
time the change has occurred. South Korea has a small number of cases, so (slightly) 256
In this work, we have attempted to quantify the level of testing which would be 258
needed to allow a country to make a near-complete return to a normal functioning of its 259
society. Among the countries considered here, we estimate that a frequency of isolating 260
individuals based on testing combined with contact-tracing raging from once every 6.2 261
days (Italy) to once every 3.5 days (U.S.) would work to keep the pandemic under a 262
“slow-burn” control. These estimates assume a test with sensitivity at or near 100% and 263
immediate isolation once a subject tests positive. Though reaching these targets would 264
necessitate an undeniably large logistic effort, home-test kits availability4 combined 265
5
with further advances in mobile device-based contact tracing can make these strategies 266
4 A recent discussion on home-test kits this can be found in “COVID-19: A cheap, simple way to control
July 2020 with Ontario being first province to test it on a larger scale: https://www.cp24.com/news/
Fig 11. Predicted daily cases in the U.S. under different rates of isolation due to
testing, or testing plus contact-tracing, accompanied by a cessation of distancing
measures.
possible. 267
disease transmission model. Although we have taken the proportion of asymptomatic 270
COVID-19 cases to be 50%, informed by a recent CDC report, assuming asymptomatic 271
infection confers immunity, this would mean a smaller remaining pool of susceptibles 272
and thus a lower current effective reproduction number. Estimates of R0 from time 273
series data of cases depend, as always, on the assumed latent and infectious periods. As 274
we have demonstrated through (Fig 7), if these periods are longer than the isolation 275
time, then it is the latter which principally drives the disease dynamics. Thus, our 276
findings about threshold isolation times are relatively robust against the possibility of a 277
6 Conclusion 279
Testing and tracing policy directions must be strongly dependant on public cooperation 280
and compliance. Populations will become anxious to resume more normal work, school 281
and social schedules, while compliance with measures will become harder to enforce. 282
covid-19-alert-app-starts-beta-testing-after-three-week-delay-1.5036434
The population must be relied on to comply with self-isolation if testing positive for the 283
virus, as well as self-isolation upon being exposed to an infected person. Last, 284
sustainable supply chains, accuracy and reliability of possible tests as well as privacy 285
issues around electronic contact tracing technology all present important, though not 286
insurmountable, hurdles countries must solve. Absent universal availability of effective 287
vaccines and treatments, the testing and tracing policies together with NPI measures are 288
much more desirable and should be the one to strive for in the immediate short-term. 289
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A Appendix 350
We want to find a relation between the exponential growth of the infected compartment 353
in an SEIRL model (10) and the reproductive number R0 around a disease-free 354
equilibrium of the type (s(0) = s̃ ≤ 1, 0, 0, 0) which arises as a possibility in a first wave 355
(s̃ = 1) or a second wave of a pandemic such as COVID-19 (s̃ < 1). 356
In this case, we conduct a similar computation as in [12], but considering the 4 357
dimensional system of equations for s, e, i, l leads us to the Jacobian of the SEIRL: 358
−βi 0 −βs 0
βi −(σ + κ1 ) βs 0
J = .
0 σ −(γ + κ) 0
0 0 κ 0
0 0 −βs̃ 0
0 −(σ + k1 ) βs̃ 0
J = .
0 σ −(γ + κ) 0
0 0 κ + κ1 0
Again we note that the linearized equations for s and l are decoupled from the equations 360
of e and i, thus, to get information on the growth rate of the infected compartment, let 361
us try to solve the linearized reduced system in (e, i) based on the reduced Jacobian: 362
−(σ + κ1 ) βs̃ Solve:
Jreduced = =⇒ det(ρI2 − Jreduced ) = 0
σ −(γ + k)
−(σ + κ1 + γ + κ)
ρ1,2 = ±
2
p 365
p
−(σ + κ1 + γ + κ) ± ((σ + κ1 ) − (γ + κ))2 + 4σβs̃
ρ1,2 = (15)
2
We first note that ((σ + κ1 ) − (γ + κ))2 + 4σβs̃ > 0, given all parameters are 366
positive. This implies that ρ1 6= ρ2 ∈ R and clearly ρ2 < 0. We check whether ρ1 > 0 by 367
looking at 368
p
((σ + κ1 ) − (γ + κ))2 + 4λβs̃ > σ + κ1 + γ + κ ⇔
(σ + κ1 )(γ + κ)
σβs̃ > (σ + κ1 )(γ + κ) ⇔ βs̃ >
σβ
(σ + κ1 )(γ + κ)
ρ1 > 0 as long as s̃ > (16)
σβ
Inequality (16) simply shows that in order to not have an exponential growth from 371
our disease free equilibrium (in other words the infection dies out), we need to allow 372
(σ + κ1 )(γ + κ)
s̃ ≤
σβ
We note that β and γ are disease-dependent values on which we cannot exert control. 374
However, κ and κ1 are parameters on which we can exert an exogenous control 375
(specifically to increase them, thus raising the upper bound on fractions s̃ with no 376
exponential growth in infected) which will be outlined in detail in the next section. 377
get: 379
p
2ρ1 + (σ + κ1 ) + (γ + κ) = ((σ + κ1 ) − (γ + κ))2 + 4σβs̃ =⇒
2 2
(σ + κ1 ) − 2(σ + κ1 )(γ + κ) + (γ + κ) + 4σβs̃ ⇐⇒
ρ21 + (σ + κ1 )(γ + κ) + ρ1 (σ + κ1 ) + ρ1 (γ + κ)
= β ⇐⇒
σs̃
382
(σ + κ1 + ρ1 )(γ + κ + ρ1 )
=β
σs̃
Following [23], we can use the next generation matrix to deduce R0 as the dominant 383
βσs̃ βs̃
(σ+κ1 )(γ+κ) γ+κ βσs̃
F V −1 = =⇒ R0 =
0 0 (σ + κ1 )(γ + κ)
Now using the expression of β in that of R0 we are able to express the effective 385
σs̃ (σ + κ1 + ρ1 )(γ + κ + ρ1 )
R0 = =⇒
(σ + κ1 )(γ + κ) σs̃
387
(σ + κ1 + ρ1 )(γ + κ + ρ1 )
R0 = and Ref f = s(t)R0 , ∀t > 0
(σ + κ1 )(γ + κ)
(σ + κ1 + ρ1 )(γ + κ + ρ1 )
R0SEIRL =
(γ + κ)σ
Let us now note that we have shown that the exponential growth factor (15), as well 390
as the R0SEIRL , are dependent on the rates κ and κ1 , that is to say, we denote by 391
p
−(σ + κ1 + γ + κ) + ((σ + κ1 ) − (γ + κ))2 + 4σβs̃
ρ(κ, κ1 ) = ,
2
392
(σ + κ1 + ρ1 )(γ + κ + ρ1 )
and by R0 (κ, κ1 ) =
(γ + κ)σ
1
Let us express the reproductive number, in general, as a function of Tlat = σ and 394
1 1
Tser = Tinf + Tlat = = Tser − Tlat =⇒ γ = . 395
γ Tser − Tlat
1 1
(ρ + )(ρ + )
(ρ + γ)(ρ + σ) Tlat Tser − Tlat
R0 = =
γσ 1
Tlat (Tser − Tlat )
397
= (ρTlat + 1)(ρ(Tser − Tlat ) + 1).
Then 398
dR
= −ρ(Tlat ρ + 1) − (Tlat ρ − Tser ρ − 1)ρ
dTlat
where we can solve for a Tlat value which maximizes R0 , namely 400
dR Tser
= 0 ⇐⇒ −2Tlat ρ + Tser ρ =⇒ Tlat = .
dTlat 2