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Introduction To Infectious Disease Modelling (EC06)
Introduction To Infectious Disease Modelling (EC06)
Modelling (EC06)
Module: EPM301 Epidemiology of Communicable Diseases
Course: PG Diploma/ MSc Epidemiology
This document contains a copy of the study material located within the computer
assisted learning (CAL) session. The first three columns designate which page,
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to refer back to specific sessions.
These study materials have been prepared by the London School of Hygiene & Tropical Medicine as part of
the PG Diploma/MSc Epidemiology distance learning course. This material is not licensed either for resale
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London School of Hygiene & Tropical Medicine September 2013 v1.0
Section 2: Introduction
Mathematical models are increasingly being used to interpret and predict the
dynamics and control of infectious diseases.
Applications include predicting the impact of vaccination strategies against common
infections such as measles and rubella, and determining optimal control strategies
against influenza, HIV and vectorborne diseases.
This session illustrates some of the areas of application of modelling and will
introduce you to some of the methods for setting up models.
Mathematical models
Physical models
The classic example of the physical or mechanical model is the ReedFrost
teaching model, developed at Johns Hopkins during the 1930s to teach medical
students about the numbers of cases which are likely to be seen over time,
following the introduction of infectious cases into a population.
Interaction: Button: Reference (pop up box appears):
Fine (1977) A commentary on the mechanical analogue to the Reed-Frost epidemic
model. Am J Epidemiol, 1977, 106(2): 87-100
It is named after two famous infectious disease epidemiologists: Reed and Wade
Hampton Frost who developed the model.
Nowadays, this whole process of simulating an epidemic can be carried out using
mathematical equations, implemented on a computer, within a matter of seconds.
Ct + 1 = St Ct r
This particular model is based on reasoning by Hamer in 1906, and was designed
to interpret the biennial cycles in measles incidence which are seen in most
populations.
Interaction: Hyperlink: seen in most populations
Output: [pop up]:
Weekly measles notifications in England and Wales, 195079 (Fine and
Clarkson, 1982)
This idea was first conceived by Ronald Ross in 1908, who reasoned that to control
malaria transmission, the density of mosquitoes in the population needed to be
reduced to a sufficiently low level.
Interaction: button: References (1):
Output:
Ross (1911) The Prevention of Malaria, 2nd ed., Murray, London.
End interaction.
This theme was subsequently developed by Kermack and McKendrick in 1927, and
then by Macdonald in 1957, who defined the basic reproduction rate as the average
number of secondary cases resulting from each infectious case when introduced into
a totally susceptible population.
Interaction: button: References (2):
Output:
Kermack WO and McKendrick AG (1927). Contributions to the mathematical theory
of epidemics. I Proceedings of the Royal Society of Medicine, 115A: 700721.
MacDonald G (1957). The Control of Epidemiology and Control of Malaria. Oxford
University Press, London.
Everyone is as likely to contact, e.g. people of different ages, social groups, etc., as
they are to contact people of their own age, social group, etc. This is also referred
to as homogenous mixing.
End interaction.
See the tabs to revise these methods.
Calculating the R0
Interaction: tabs:
Tab 1:
Revision I
As you saw in EC03, if a population has a rectangular
Interaction: hyperlink: rectangular
Output:
End interaction.
age distribution (also known as a Type I distribution), such as that seen in many
Western populations, R0 can be calculated using the expression R0 = L/A
Here, L is the life expectancy and A is the average age at infection.
Tab 2:
Revision II
The expression for the R0 for a population with a negative exponential
End interaction.
age distribution (also known as a Type II distribution), such as that seen in some
developing countries is analogous to that for a population with a rectangular age
distribution:
R0 = 1 + L/A
Here, L is the average life expectancy and A is the average age at infection. See
Anderson and May (1991) for the derivation of this expression.
Interaction: Button: "References":
Anderson and May (1991)
Infectious Disease of Humans.
Dynamics and Control. Oxford
University Press. Oxford
Question:
If vaccination is available only for children, is the level of coverage required to
control transmission in population A higher, equal to or lower than that in population
B?
Interaction: hotspots:
Hotspot 1:
Higher
Output [correct response]:
Correct
Control is easiest to achieve if it is targeted at the individuals who are the most
active transmitters. In population A, young individuals are responsible for a smaller
proportion of all the transmission events (2/8) than in population B (4/8). As a
Given that the true degree of contact between different population groups (e.g. the
young and the old) is poorly understood, in some countries e.g. the UK, models are
used to explore the effect of several different assumptions about contact between
individuals on the impact of vaccination against measles, pneumococcal,
meningococcal and other infections.
Interaction: tabs
Tab 1:
Example
In the UK, uptake of MMR vaccination has been variable (and sometimes below the
herd immunity threshold for measles), as a result of public concerns over its safety.
Changes in the proportion of individuals who are susceptible to measles infection, as
a result of MMR vaccination and ongoing transmission are used to calculate the
overall net reproduction number for different assumptions about contact between
individuals.
Interaction: Button: "References":
Gay NJ, Hesketh LM,
Morgan-Capner P, Miller E (1995)
Interpretation of serological
surveillance data for measles
using mathematical models:
implications for vaccine strategy.
Epidemiol Infect. 1995;
115(1): 139-56.
Tab 2:
Example (cont)
The size of the net reproduction number then provides an indication of the chance of
an outbreak of measles; models incorporating different assumptions about contact
can provide insight into the numbers of cases which might occur in different age
groups.
If no cases have been observed e.g. for 1 year, what is the probability that control
has been achieved?
These questions have been explored in relation to rubella, measles, and more
recently polio.
Interaction: Button: References
Output:
Anderson RM and Grenfell BT (1986). Quantitative investigation of different
vaccination policies for the control of congenital rubella syndrome (CRS) in the
UK. J Hyg. Camb, 96, 30533.
Anderson RM and May RM (1985). Agerelated changes in the rate of disease
transmission: implications for the design of vaccination programmes. J Hyg.
Camb, 94, 365436.
Eichner M, Hadeler KP, Ditz K. Stochastic models for the eradication of
poliomyelitis: minimum population size for polio virus persistence. Model for
Infectious Diseases: Their Structure and Relation to Data. Eds. V Isham, G
Medley. Cambridge University Press, 1996.
Trotter CL, Gay NJ, Edmunds WJ (2005). Dynamic models of meningococcal
carriage, disease, and the impact of serogroup C conjugate vaccination. Am J
Epidemiol. 2005, July 1; 162(1):89100.
End interaction.
To illustrate this use of modelling, we shall focus on one specific example, namely
the relationship between rubella, the incidence of Congenital Rubella Syndrome
and rubella vaccination coverage.
Question: Assuming that neither country had introduced rubella or MMR vaccination
during the 1980s, in which country would you have expected to see a higher
incidence (per live birth) of Congenital Rubella Syndrome ?
Interaction: Buttons:
Button 1: UK
Output:
UK
Correct. The proportion of women of childbearing age who have antibodies (i.e.
immunity) to rubella is higher in China than in the UK. You might therefore expect
the number of new infections per capita among women in China to be lower than
that in the UK, and for the incidence of CRS per 1000 live births to be
correspondingly lower in China.
Button 2: China
Output:
China
Thats not quite right. The proportion of women of childbearing age who have
antibodies (i.e. immunity) to rubella is higher in China than in the UK. You might
therefore expect the number of new infections per capita among women in China to
be lower than that in the UK, and for the incidence of CRS per 1000 live births to be
correspondingly lower in China.
Button 3: References
Output:
Farrington CP (1990) Modelling forces of infection for measles, mumps and rubella.
Statistics in Medicine, 9, 95367.
Wannian (1985). Rubella in the Peoples Republic of China. Rev Inf Dis; 7 (S72).
In China, the introduction of infant MMR vaccination at below the herd immunity
threshold might therefore result in the emergence of a new problem of CRS and,
potentially, a greater relative increase in the burden of CRS than in the UK.
risk of infection
The introduction of infant vaccination will lead to an increase in the proportion of
women of childbearing age who are susceptible.
On the other hand, it will lead to a reduction in the risk of infection.
The net effect on the incidence of CRS will depend on whether the increase in the
proportion of women of childbearing age who are susceptible outweighs the
reduction in the risk of infection
Typed out by M. A.
Predictions of the ratio between the numbers of cases of CRS after the
introduction of vaccination among infants at different levels of coverage (Xaxis) and that before the introduction of vaccination (Anderson and May
(1983)
Proportion vaccinated
End interaction.
The different lines are predictions for settings in which the average age at
infection before the introduction of vaccination ranged between 3 and 18
years.
Interaction: button: References
Output:
These were attributed to improvements in hygiene and hence a reduction in the risk
of infection and an increasing proportion of individuals being infected with polio virus
for the first time when adults. (Note that infection during adulthood carries a greater
risk of paralytic polio than does infection as a child).
Interaction: button: references
Output:
Anderson and May (1991). Infectious Diesases of Humans: Dynamics and Control.
Oxford University Press, Oxford.
Mumps outbreaks of mumps have been seen among adolescents and young
adults in the UK and other Western countries since the year 2000.
This has been attributed to the introduction of MMR vaccination among young
children, leading to a reduced risk of infection in the overall population, with
individuals born before the introduction of vaccination reaching adult life still
susceptible to infection.
Interaction: button: references
Output:
Savage E, Ramsay M, White J, Beard S, Lawson H, Hunjan R, Brown D. Mumps
outbreaks across England and Wales in 2004: Observational study. BMJ,
2005;330(7500):111920.
shows predictions of the Hamer model described earlier, adapted to take daily time
steps, rather than time steps of 1 serial interval.
Note that this model, which is more realistic than the Hamer model, predicts that the
cycles should ultimately damp out. The fact that this damping out is inconsistent
with the observed data, suggests that extrinsic factors, which had not been
incorporated in the model, help to sustain the cycles.
Tab 2:
The diagram
Interaction: hyperlink: diagram
Output:
We will first describe what we mean by deterministic models and return to the other
kinds of models (stochastic models) later in this session.
For immunizing infections, for example, it would be useful to stratify people into
those who are Susceptible, Infected (but not yet infectious), Infectious and
Immune, as follows:
Note that the InfEcted class is also sometimes referred to as the _xposed_class;
this however, can be confusing since everyone can be considered to be exposed.
Returning to our measles model, and considering the susceptible individuals, the
equations need to specify the fact that:
Number susceptible at time t + 1 =
Number susceptible at time t
Output:
For example, if 10% (= t) of susceptible individuals are newly infected between
today and tomorrow, and we have 500 (St) susceptible individuals present today,
then the number of individuals who are newly infected between today and
tomorrow is 0.1 * 500 = 50.
We can now formally write down the equations for the number of susceptible
individuals at time t + 1 as follows:
St + 1 = St tSt
or as
St + 1 = St StIt
Returning to our model diagram and now considering the individuals in the
infEcted category, we see that
the equations need to specify the fact that:
Number of individuals in the infEcted category at time t + 1
=
Number of individuals in the infEcted category at time t
+
Number of individuals who are newly infEcted between time t and t + 1
Et + 1 = Et + StIt f Et
Here we are assuming that the proportion of infected individuals who develop
infectious disease in each time step is the same over time we will return to this
assumption later.
Returning to our model diagram and now considering the individuals in the
infectious category, we see that the equations need to specify the fact that:
Number of individuals who are infectious at time t + 1
=
Number of individuals who were infectious at time t
+
Number of individuals who develop infectious disease between time t and
t + 1.
Number of individuals who recover from infectious disease between time t and t + 1
The number of individuals who recover from infectious disease between time t and
t + 1, in turn, is just the product of the proportion of infectious individuals who
recover from infectious disease between time t and t + 1 (we will denote this
proportion by r for now) and the number of infectious individuals at time t. i.e. r
It
We will return to the definition and derivation of r later in this session.
It + 1 = It + fEt rIt
Exercise:
Write down the equation for the number of immune individuals at time t + 1 (Rt +
1) in terms of Rt, It and r, and check your answer by clicking on the answer box.
Interaction: Button: Answer:
Answer
The equations need to specify the fact that:
Number of individuals who are immune at time t + 1 =
Number of individuals who were immune at time t
+
Number of individuals who recover from infectious disease between time t and t+1
Using mathematical notation, this equation can be written as:
Rt+1 = Rt + rIt
or equivalently
St + 1 = St tSt
Et + 1 = Et + tSt fEt
It + 1 = It + fEt r It
Rt + 1 = Rt + r It
At present, this model does not incorporate births entering the population or
individuals dying.
To incorporate births into the population, we would need to change the equation
for the number of susceptible individuals (assuming, for now, that individuals are
born susceptible as follows:
St + 1 = St St It + Bt
where Bt is the number of individuals born between time t and t + 1.
This is a useful check to see whether your equations have any errors.
End interaction.
then the risk is approximately equal to the rate.
step , the model can be set up in a spreadsheet to predict the total number of
individuals present in each of the compartments over time.
We shall now illustrate how this is done in further detail.
Hyperlink 2:
Check Excel expression for immune individuals for day 1
Output:
= E44=D44*rec_rate
Compare your graph of the number of susceptible, infectious and immune
individuals to the graph here.
Interaction: hyperlink: graph here
Output:
If the two differ, refer to spreadsheet measles1solna.xls to check that you have
typed in the equations correctly.
The graph shows the classic epidemic curve, i.e. after a peak in the incidence of
infectious cases, no further cases arise in the population.
This is attributable to the depletion of susceptible individuals: all have been infected,
have developed disease and are immune to further infection.
individuals on day 1 to incorporate births into and deaths out of the population
(assuming, for now, that individuals are born susceptible.
Step 5: Copy all the equations for day 1 down until day 25550 (i.e. 70 years).
Interaction: button: Example
Output:
Check Excel expressions for day 1:
Susceptible individuals:
= B44+beta *B44*D44+num_births-B44*m_rate
Infected individuals:
= C44+beta *B44*D44-dis_rate*C44-C44*m_rate [Note: yes, it is C44-C44]
Infectious individuals:
= D44+C44*dis_rate-D44*rec_rate-D44*m_rate
Immune individuals
=E44+D44*rec_rate-E44*m_rate
Step 6: Select rows 18 and 37 together, click with your right mouse button and
choose the unhide option.
NB To select two rows together, click on the grey row header for the first row that
you're interested in, and either hold down the left mouse button and drag the
mouse down to the grey row header for the next row, OR hold the shift key down
and click on the grey row header for the second row.
You should now see the following graph
Interaction: hyperlink: following graph
Output:
of the numbers of susceptible and immune individuals over time. If your graph
differs from this graph, check your equations against the equations in
measles1solnb.xls.
The graph shows that the numbers of susceptible and immune individuals cycle over
time.
Step 7: To see this, double click with your mouse on the righthand yaxis in the
figure showing the longterm trends in the spreadsheet and change the scale to go
from zero to 20.
We shall discuss the relationship between the peaks in the disease incidence and
the numbers of susceptible individuals, after we have considered the derivation of
the input parameters in the model.
Before continuing, please save your spreadsheet as measlesfin1.xls
Question:
Which of the bars in the above figure are likely to reflect the seroprevalence among
females in rural areas?
Interaction: Buttons:
Button 1: White Bars
Output:
White Bars
Correct. For each age group, the seropositivity is lower among individuals reflected
in the white bars than for the individuals reflected in the black bars, suggesting that
these individuals face a correspondingly lower risk of infection, i.e. they probably live
in rural areas.
Button 2: Black Bars
Output:
Black Bars
Thats not quite right. For each age group, the seropositivity is lower among
individuals reflected in the white bars than for the individuals reflected in the black
bars, suggesting that these individuals face a correspondingly lower risk of infection,
i.e. they probably live in rural areas.
Button 3: References
Output:
Dowdle WR, Ferrera W, De Salles Gomes LF, King D, Kourany M, Madalengoitia J,
Pearson E, Swanston WH, Tosi HC, Vilches AM. WHO collaborative study on the
seroepidemiology of rubella in Carribbean and Middle and South American
populations in 1968. Bull World Health Organ. 1970; 42(3):41922.
individuals living in many Western countries in the year 1900 probably contacted
more individuals than their counterparts who are alive today.
Interaction: Button: References:
Hunt S. Housing-related
disorders. In: Charlton J, Murphy
M (eds.), The health of adult
Britain, 1841-1994. Volume 1;
chap. 10, 157-70. The Stationery
Office; London 1997.
may also change over time as a result of changes in behaviour or interventions.
During past influenza pandemics, for example, the amount of contact between
individuals changed over time, e.g. with theatres and concert halls shutting and
office closing hours being staggered to avoid the congregation of individuals.
is difficult to measure directly for the majority of infections, excepting possibly,
those involving a vector or sexual transmission. Consequently, it is typically
calculated using indirect methods involving analysis of seroprevalence data and/or
the basic reproduction number.
Considering population of a given size, is the value for greater for measles or for
rubella?
Interaction: button: Measles
Output:
Measles
Correct. In a population comprising 100 individuals, the value for is 0.019 per
day for measles and 0.0064 per day for rubella.
Interaction: button: Rubella
Output:
Rubella
Thats not quite right. In a population comprising 100 individuals, the value for is
0.019 per day for measles and 0.0064 per day for rubella, i.e. it is higher for
measles than for rubella.
For example, the average infectious period for measles is 7 days; the rate at which
infectious individuals recover from infectious disease is 1/7 = 0.143 per day.
End interaction.
Distribution of the time interval between infection and onset of
infectiousness, assuming that individuals develop infectious disease at a
constant rate of 0.143/day
Days since infection
It is possible to refine this assumption e.g. to assume that the rate of infectious
disease onset depends on time since infection.
In practice, the level of complexity incorporated in the model usually depends on
the kind of question asked.
You should also notice that, as for measles, all individuals are immune in the
population by the end of the first 200 days following the introduction of the infectious
case.
Interaction: button: Graph
Output:
If Rn is bigger than one (i.e. each infectious case is leading to more than one
secondary infectious case), then the disease incidence should increase.
If Rn is less than one (i.e. each infectious case is leading to less than one secondary
infectious case), then the disease incidence should decrease.
We will now use our spreadsheet model to explore this relationship further.
Step 1: Open up the spreadsheet measles2.xls. It is very closely related to the
spreadsheet you have just been using, except that we now have some cells set up
for the proportion of susceptible and immune individuals in the population for day
1.
Step 2. Set up an appropriate expression for the net reproduction number on day 1
and copy this expression down until the 18250th day.
Interaction: Button: Hint:
The expression for the net reproduction number on day 1 should be: =G81*R0
(end interaction)
which plots the Rn on the left hand yaxis and the disease incidence on the right
hand yaxis in the population. The xaxis goes from the 14600th day (ie the 40th
year) to the 18250th day (ie the 50th year). The gridlines on the xaxis occur at 2
year intervals.
(If Figure 1 in your spreadsheet does not resemble this Figure, you may like to check
the equations in your spreadsheet against the equations in measles2ansa.xls).
Question: What is the value of the Rn when the disease incidence peaks or reaches
a trough?
Interaction: hotspots (set 1):
Hotspot 1: ~>1
Thats not quite right. At the peak or trough of the disease incidence, Rn is
approximately equal to 1.
Hotspot 2: ~<1
Thats not quite right. At the peak or trough of the disease incidence, Rn is
approximately equal to 1.
Hotspot 3: =1
Correct. At the peak or trough of the disease incidence, Rn is approximately equal
to 1.
End hotspots (set 1).
Question: What is happening to the disease incidence when Rn~<1?
Interaction: hotspots (set 2):
Hotspot 1: Increasing
Thats not quite right. When Rn~<1, the disease incidence is increasing.
Hotspot 2: Decreasing
Correct. When Rn~<1, the disease incidence is increasing.
Hotspot 3: Unchanging
Thats not quite right. When Rn~<1, the disease incidence is increasing.
Question: What is happening to the disease incidence when Rn~>1?
Interaction: hotspots (set 3)
Hotspot 1: Increasing
Correct. When Rn~>1, the disease incidence is increasing.
Hotspot 2: Decreasing
Thats not quite right. When Rn~>1, the disease incidence is increasing.
Hotspot 3: Unchanging
Thats not quite right. When Rn~>1, the disease incidence is increasing.
(end interaction)
which plots the proportion susceptible on the left hand yaxis and the disease
incidence on the right hand yaxis.
Question. What proportion of the population is susceptible to infection when the
disease incidence peaks or troughs?
Interaction: button: Answer
Output:
At a peak or trough in disease incidence, the proportion of individuals which are
susceptible is approximately 0.077.
You should remember from previous sessions that, for transmission of an infectious
disease to persist in a population, the proportion of the population which is immune
to infection has to be below the herd immunity threshold(HIT), which is given by:
HIT = 1 1/R0
Question: According to this expression, what should be the herd immunity threshold
for the infection in this model?
Interaction: button: Answer
Output:
The R0 is 13, so the herd immunity threshold is 11/R0 which is approximately
equal to 92.3%.
(end interaction)
which plots the proportion immune in the population together with the disease
incidence.
Question: What proportion of the population is immune when the disease
incidence peaks or troughs?
Interaction: button: Answer
Output:
At a peak or a trough in the disease incidence, the proportion of the population
which is immune is approximately 0.923, i.e. equal to the herd immunity threshold.
(Note that this is equal to 1 0.077, the proportion of the population which is
susceptible at this time, as we saw earlier on EC06p15c3)
End interaction.
Question: What do you notice about the proportion of the population which is
immune when the disease incidence is a) declining? b) increasing?
Interaction: button: Answer A
Output:
When the disease incidence is declining, the proportion of the population which is
immune is always above 0.923, i.e. above the herd immunity threshold.
End Answer A interaction.
Interaction: button: Answer B
Output:
When the disease incidence is increasing, the proportion of the population which is
immune is always below 0.923, i.e. above the herd immunity threshold.
The exercise also highlights the fact that cycles in the incidence of an immunizing
infection occur because of cycles in the proportion of individuals who are susceptible
and immune.
who are susceptible i.e. more susceptibles are being removed than are being added
to the population. (point C)
Once the proportion of susceptible individuals has decreased to be below 1/R0, each
case starts to lead to less than one secondary case and the disease incidence
decreases (point D).
Interaction: button: Graph
Output:
T 2
L( D + D' )
R 0 1
Here, L is the average life expectancy, and D and D are the durations of the latent
and infectious periods respectively. is the universal constant, given approximately
by 3.14.
Use this formula to calculate the interepidemic period in a population in which the
life expectancy is 60 years for
a) measles (R0=13, D=8 days, D=7 days)
b) rubella (R0=7, D=10 days, D=11 days)
T 2 x3.14
60 x365(7 + 8)
= 1039days = 2.85 years
13 1
T 2 x3.14
60 x365(11 + 10)
= 1739days = 4.76 years
7 1
Step 1: Change the size of the time step (in cell F4) to be 0.5, 2, 3, 4 and 5 days
and look at predictions of the longterm disease incidence.
for the measles model we were discussing earlier differs from that for the
difference equations.
Interaction: hyperlink: difference equations
Output:
These differences are discussed in the optional reading section for this session.
If a stochastic model is run many (e.g. 1000) times, then the average result from all
of these runs should be very similar to the outcome from a deterministic model.
Output:
Ferguson NM, Cummings DA, Cauchemez S, Fraser C, Riley S, Meeyai A,
Iamsirlthaworn S, Burke DS. Strategies for containing and emerging influenza
pandemic in Southeast Asia. Nature. 2005 Sep 8;437(7056):20914.
End interaction.
The main drawback of complex models is that, because they incorporate many
different parameters and assumptions, the individual effect of a particular factor on
the model prediction becomes difficult to understand.
19.1: Summary
Models are classified into those which are deterministic and those which are
stochastic.
Deterministic models describe what happens on average in a population.
Stochastic models incorporate chance variation in e.g. the number of individuals
infected and might be used to address questions such as:
what is the probability of an outbreak following the introduction of one case into a
given population?
19.2: Summary
Deterministic models can be set up using either difference equations of differential
equations.
Difference equations describe the transmission dynamics of an infection using
discrete time steps of e.g. 1, 2 days etc.
Differential equations describe the transmission dynamics using continuous time.
Interaction: tabs:
Tab 1: Difference Equations
An example of a model for an immunizing infection set up using difference
equations:
19.3: Summary
The most important parameter in infectious disease models is , defined as the
probability of an effective contact between two specific individuals per unit time.
For an immunizing infection in a randomly mixing population, it can be calculated
using the expression:
R0/(ND)
where N is the population size and D is the duration of infectiousness.
For simple models, the transition rates, (e.g. rates of infectious disease onset, the
recovery rate) can be calculated using the expressions:
1/(average latent period)
or
1/(average duration of infectiousness)
assuming that these rates are constant over time.
19.4: Summary
The level of complexity incorporated in a model depends on the question being
addressed.
For example, a simple model of the transmission dynamics of measles following the
introduction of 1 infectious case into a totally susceptible population can provide
useful insight into the factors underlying the cycles in incidence e.g. the basic
reproduction number.
On the other hand, such a model would need to be extended to make predictions
about the numbers of cases in specific age groups, or the effect of nonrandom
mixing on the impact of control strategies.
Returning to the above model and considering the susceptible category, for
example, as noone enters the susceptible class, the rate of change in the
number of susceptible individuals is given by the expression:
the number of susceptible individuals who become newly infected per unit
time
Using mathematical notation, this would be written as:
dS(t)
= -S(t)I(t)
dt
Considering the infected category in the same model, newly infected individuals
enter this category and diseased individuals exit this category.
The rate of change in the number of infected individuals is given by the expression:
+ the number of susceptible individuals who become newly infected per unit time
the number of infected individuals who develop infectious disease per unit time.
Using mathematical notation, the rate of change in the number of infected
individuals would be written as:
dE(t)
= -S(t)I(t)-fE(t)
dt
Exercise:
Considering the above model, write down the differential equations for the rate of
change in the number of
Interaction: Button: (a)infectious individuals
Output:
A total of fE(t) Infection individuals enter this category per unit time and rI(t) exit
this category per unit time. The rate of change in the number of infectious
individuals is therefore given by:
dl (t )
= fE (t ) rl (t )
dt
Interaction: Button: (b) immune individuals
Output:
A total of rI(t) Infectious individuals enter this category per unit time and no
individuals exit this category per unit time. The rate of change in the number of
immune individuals is therefore given by:
dR(t )
= rl (t )
dt
Exercise:
Write down the differential equations corresponding to the following model
diagram, assuming that no individuals are born into or die from the population.