Eric Forgoston1, Lora Billings, Ira B Schwartz. 1. Nonlinear Dynamical Systems Section, Plasma Physics Division, US Naval Research Laboratory, Code 6792, Washington, DC 20375, USA. eric.forgoston.ctr@nrl.navy.mil
Abstract
We consider a stochastic susceptible-exposed-infected-recovered (SEIR) epidemiological model. Through the use of a normal form coordinate transform, we are able to analytically derive the stochastic center manifold along with the associated, reduced set of stochastic evolution equations. The transformation correctly projects both the dynamics and the noise onto the center manifold. Therefore, the solution of this reduced stochastic dynamical system yields excellent agreement, both in amplitude and phase, with the solution of the original stochastic system for a temporal scale that is orders of magnitude longer than the typical relaxation time. This new method allows for improved time series prediction of the number of infectious cases when modeling the spread of disease in a population. Numerical solutions of the fluctuations of the SEIR model are considered in the infinite population limit using a Langevin equation approach, as well as in a finite population simulated as a Markov process.
We consider a stochastic susceptible-exposed-infected-recovered (SEIR) epidemiological model. Through the use of a normal form coordinate transform, we are able to analytically derive the stochastic center manifold along with the associated, reduced set of stochastic evolution equations. The transformation correctly projects both the dynamics and the noise onto the center manifold. Therefore, the solution of this reduced stochastic dynamical system yields excellent agreement, both in amplitude and phase, with the solution of the original stochastic system for a temporal scale that is orders of magnitude longer than the typical relaxation time. This new method allows for improved time series prediction of the number of infectious cases when modeling the spread of disease in a population. Numerical solutions of the fluctuations of the SEIR model are considered in the infinite population limit using a Langevin equation approach, as well as in a finite population simulated as a Markov process.
The reduction in high-dimensional, stochastic systems is an
important and fundamental problem in nonlinear dynamical systems. In this article, we present
a general theory of stochastic model reduction, which is based on a normal form coordinate
transform. This nonlinear, stochastic projection allows for the deterministic and stochastic
dynamics to interact correctly on the lower-dimensional manifold so that the dynamics
predicted by the reduced, stochastic system agrees well with the dynamics predicted by the
original, high-dimensional stochastic system. Although the method may be applied to any
physical or biological system with well-separated time scales, here we apply the method to an
epidemiological model. We show that when compared with the original stochastic epidemic model,
the reduced model properly captures the initial and recurrent disease outbreaks, both in
amplitude and phase. This long-term accuracy of the reduced model allows for the application
of effective disease control where phase differences between outbreak times and vaccine
controls are important. Additionally, in practice, one can only measure the number of
infectious individuals in a population. Our method allows one to predict the number of
unobserved exposed individuals based on the observed number of infectious individuals.
INTRODUCTION
The interaction between deterministic and stochastic effects in population dynamics has
played, and continues to play, an important role in the modeling of infectious diseases. The
mechanistic modeling side of population dynamics is well known and established. These models typically are assumed to be
useful for infinitely large, homogeneous populations, and arise from the mean field analysis
of probabilistic models. On the other hand, when one considers finite populations, random
interactions give rise to internal noise effects, which may introduce new dynamics.
Stochastic effects are quite prominent in finite populations, which can range from
ecological dynamics to childhood epidemics
in cities. For homogeneous
populations with seasonal forcing, noise also comes into play in the prediction of large
outbreaks. Specifically,
external random perturbations change the probabilistic prediction of epidemic outbreaks as
well as its control.When geometric structure is applied to the population, the interactions are modeled as a
network. Many types of static
networks which support epidemics have been considered. Some examples include small world
networks, hierarchical networks, and transportation networks of patch
models. In addition, the fluctuation
of epidemics on adaptive networks, where the wiring between nodes changes in response to the
node information, has been examined. In
adaptive network models, even the mean field can be high dimensional, since nodes and links
evolve in time and must be approximated as an additional set of ordinary differential
equations.Another aspect of epidemic models, which is often of interest, involves the inclusion of a
time delay. The delay term makes the analysis significantly more complicated. However, it is
possible to approximate the delay by creating a cascade consisting of a large number of
compartments. For example, one could
simulate the delay associated with a disease exposure time with several hundred “exposed”
compartments.These model examples are just a few of the types of very high-dimensional models that are
currently of interest. As a result of the high dimensionality, there is much computation
involved, and the analysis is quite difficult. In particular, real-time computation is not
currently possible. However, there are usually many time scales that are well separated (due
typically to a large range in order of magnitude of the parameters) when considering such
high-dimensional problems. In the presence of well-separated time scales, a model reduction
method is needed to examine the dynamics on a lower-dimensional space. It is known that
deterministic model reduction methods may not work well in the stochastic realm, which
includes epidemic models. The purpose of
this article is to examine a method of nonlinear, stochastic projection so that the
deterministic and stochastic dynamics interact correctly on the lower-dimensional manifold
and predict correctly the dynamics when compared with the full system. Because the noise
affects the timing of outbreaks, it is essential to produce a low-dimensional system that
captures the correct timing of the outbreaks as well as the amplitude and phase of any
recurrent behavior.We will demonstrate that our stochastic model reduction method properly captures the
initial disease outbreak and continues to accurately predict the outbreaks for time scales,
which are orders of magnitude longer than the typical relaxation time. Furthermore, in
practice, real disease data include only the number of infectious individuals. Our method
allows us to predict the number of unobserved exposed individuals based on the observed
number of infectious individuals.For stochastic model reduction, there exist several potential methods for general problems.
For a system with certain spectral requirements, the existence of a stochastic center
manifold was proven in Ref. 18. Nonrigorous
stochastic normal form analysis (which leads to the stochastic center manifold) was
performed in Refs. 19–22. Rigorous
theoretical analysis of normal form coordinate transformations for stochastic center
manifold reduction was developed in Refs. 23 and
24. Later, another method of stochastic normal form reduction was developed, in which any anticipatory convolutions
(integrals into the future of the noise processes) that appeared in the slow modes were
removed. Since this latter analysis makes the construction of the stochastic normal form
coordinate transform more transparent, we use this method to derive the reduced stochastic
center manifold equation.Figure 1 shows a schematic demonstrating our approach
to the problem. We consider a high-dimensional system along with its corresponding reduced
low-dimensional system. In this article, two types of low-dimensional system are discussed:
a reduced system based on deterministic center manifold analysis and a reduced system based
on a stochastic normal form coordinate transform. Regardless of the type of low-dimensional
system being considered, a common noise is injected into both the high-dimensional and
low-dimensional models, and an analysis of the solutions found using the high and
low-dimensional systems is performed.
FIG. 1.
Schematic demonstrating the injection of a common noise into both the high-dimensional
system and its associated low-dimensional system.
In this article, as a first study of a high-dimensional system, we consider the
susceptible-exposed-infected-recovered (SEIR) epidemiological model with stochastic forcing.
As previously mentioned, we could easily consider a SEIR-type model where the exposed class
was modeled using hundreds of compartments. Since the analysis is similar, we consider the
simpler standard SEIR model to demonstrate the power of our method. Section II provides a complete description of this model. Section
III describes how to transform the deterministic SEIR
system to a new system that satisfies the spectral requirements needed to apply the center
manifold theory. After the theory is used to find the evolution equations that describe the
dynamics on the center manifold, we show in Sec. IV how
the reduced model that is found using the deterministic result incorrectly projects the
noise onto the center manifold. Section V demonstrates
the use of a stochastic normal form coordinate transform to correctly project the noise onto
the stochastic center manifold. A discussion and the conclusions are contained,
respectively, in Secs. VI and VII.
THE SEIR MODEL FOR EPIDEMICS
We begin by describing the stochastic version of the SEIR model found in Ref. 26. We assume that a given population may be divided into
the following four classes that evolve in time:Susceptible class
consists of those individuals who may contract the
disease.Exposed class
consists of those individuals who have been infected by the disease but are not yet
infectious.Infectious class
consists of those individuals who are capable of transmitting the disease to
susceptible individuals.Recovered
class
consists of those individuals who are immune to the
disease.Furthermore, we assume that the total population size, denoted as , is
constant and can be normalized to ,
where ,
,
,
and .
Therefore, the population class variables ,
, , and
represent fractions of the total
population. The governing equations for the stochastic SEIR model arewhere
is the standard deviation of the noise intensity .
Each of the noise terms
describes a stochastic, Gaussian white force that is characterized by the correlation
functionsAdditionally, represents a constant birth and death
rate, is the contact rate,
is the rate of infection, so that
is the mean latency period, and is the rate of recovery, so that
is the mean infectious period. Although the contact rate could
be given by a time-dependent function (e.g., due to seasonal fluctuations), for simplicity,
we assume to be constant. Throughout this article,
we use the following parameter values: ,
,
,
and .
Disease parameters correspond to typical measles values. Note that any other biologically meaningful parameters may be
used as long as the basic reproductive rate .
The interpretation of
is the number of secondary cases produced by a single infectious individual in a population
of susceptibles in one infectious period.As a first approximation of stochastic effects, we have considered additive noise. This
type of noise may result from migration into and away from the population being
considered. Since it is difficult to
estimate fluctuating migration rates, it
is appropriate to treat migration as an arbitrary external noise source. Also, fluctuations
in the birth rate manifest itself as additive noise. Furthermore, as we are not interested
in extinction events in this article, it is not necessary to use multiplicative noise. In
general, for the problem considered here, it is possible that a rare event in the tail of
the noise distribution may cause one or more of the ,
, and
components of the solution to become negative. In this article, we will always assume that
the noise is sufficiently small so that a solution remains positive for a long enough time
to gather sufficient statistics. Even though it is difficult to accurately estimate the
appropriate noise level from real data, our choices of noise intensity lie within the huge
confidence intervals computed in Ref. 29. The case
for multiplicative noise will be considered in a separate paper.Although
in the deterministic system, one should note that the dynamics of the stochastic SEIR system
will not necessarily have all of the components sum to unity. However, since the noise has
zero mean, the total population will remain close to unity on average. Therefore, we assume
that the dynamics is sufficiently described by Eqs. (1a)–(1c). It should be noted that even if
for some , the noise allows for the reemergence of
the epidemic.
DETERMINISTIC CENTER MANIFOLD ANALYSIS
One way to reduce the dimension of a system of equations is through the use of
deterministic center manifold theory. In general, a nonlinear vector field can be
transformed so that the linear part (Jacobian) of the vector field has a block diagonal form
where the first matrix block has eigenvalues with positive real part, the second matrix
block has eigenvalues with negative real part, and the third matrix block has eigenvalues
with zero real part. These blocks
are associated, respectively, with the unstable eigenspace, the stable eigenspace, and the
center eigenspace. If we suppose that there are no eigenvalues with positive real part, then
orbits will rapidly decay to the center eigenspace.In order to make use of the center manifold theory, we must transform Eqs. (1a)–(1c) to a new system of equations
that has the necessary spectral structure. The theory will allow us to find an invariant
center manifold passing through the fixed point to which we can restrict the transformed
system. Details regarding the transformation can be found in Sec. III A, and the computation of the center manifold can be found in Sec.
III B.
Transformation of the SEIR model
Our analysis begins by considering the governing equations for the stochastic SEIR model
given by Eqs. (1a)–(1c). We neglect
the
terms in Eqs. (1a)–(1c) so that we
are considering the deterministic SEIR system. This deterministic system has two fixed
points. The first fixed point isand
corresponds to a disease free or extinct equilibrium state. The second fixed point
corresponds to an endemic state and is given byTo ease the analysis, we define a new set of variables, ,
, and , as
,
,
and .
These new variables are substituted into Eqs. (1a)–(1c).Then, treating as a small parameter, we rescale
time by letting .
We may then introduce the following rescaled parameters:
and ,
where
and
are .
The inclusion of the parameter as a new state
variable means that the terms in our rescaled system which contain
are now nonlinear terms. Furthermore,
the system is augmented with the auxiliary equation .
The addition of this auxiliary equation contributes an extra simple zero eigenvalue to the
system and adds one new center direction that has trivial dynamics. The shifted and
rescaled augmented system of equations iswhere
the endemic fixed point is now located at the origin.The Jacobian of Eqs. (5a)–(5d)
is computed to zeroth order in and is evaluated at
the origin. Ignoring the components, the Jacobian has only
two linearly independent eigenvectors. Therefore, the Jacobian is not diagonalizable.
However, it is possible to transform Eqs. (5a)–(5c) to a block diagonal form with the eigenvalue structure that is needed
to use the center manifold theory. We use a transformation matrix
consisting of the two
linearly independent eigenvectors of the Jacobian along with a third vector chosen to be
linearly independent. There are many choices for this third vector; our choice is
predicated on keeping the vector as simple as possible. This transformation matrix is
given asUsing
the fact that ,
then the transformation matrix leads to the following definition of new variables:
, , and
,The application of the transformation matrix to Eqs. (5a)–(5c) leads to the transformed evolution equations given
by
Center manifold equation
The Jacobian of Eqs. (8a)–(8d)
to zeroth order in and evaluated at the origin
iswhich
shows that Eqs. (8a)–(8d) may
be rewritten in the formwhere
,
,
is a constant matrix with
eigenvalues that have negative real parts, is
a constant matrix with eigenvalues that have zero real parts, and
and
are nonlinear functions in
, , and . In
particular,Therefore, the system will rapidly collapse onto a lower-dimensional manifold given by
center manifold theory. Furthermore,
we know that the center manifold is given bywhere
is an unknown function.Substitution of Eq. (14) into Eq. (8a) leads to the following center manifold
condition:In
general, it is not possible to solve the center manifold condition for the unknown
function .
Therefore, we perform the following Taylor series expansion of
in , , and
:where
,
,
,
,
are unknown coefficients that are
found by substituting the Taylor series expansion into the center manifold condition and
equating terms of the same order. By carrying out this procedure using a second-order
Taylor series expansion of , the center manifold equation
iswhere
so that provides a count of the number of
, , and
factors in any one term. Substitution
of Eq. (17) into Eqs. (8b) and (8c) leads to the following
reduced system of evolution equations, which describe the dynamics on the center
manifold
INCORRECT PROJECTION OF THE NOISE ONTO THE STOCHASTIC CENTER MANIFOLD
Transformation of the stochastic SEIR model
We now return to the stochastic SEIR system given by Eqs. (1a)–(1c). The shift in the fixed point to the origin will not
have any effect on the noise terms, so that the stochastic version of the shifted
equations isAs Eqs. (19a)–(19c) are
transformed using Eqs. (7a)–(7c),
the
scaling, the
scaling, and the
time scaling, the noise also is scaled so that the stochastic transformed equations are
given bywhereThe
stochastic terms ,
,
and
in Eqs. (20a)–(20c) are still
additive Gaussian noise processes. However, Eqs. (21a)–(21c) show how the transformation has acted on the
original stochastic terms ,
,
and
to create new noise processes, which have a variance different from that of the original
noise processes. Also note that we have suppressed the argument of
,
,
and
in Eqs. (20a)–(20c). The time
scaling means that these noise terms should be evaluated at .The system of equations given by Eqs. (20a)
and (21c) is an exact transformation of the system of equations given by Eqs.
(1a)–(1c). We numerically
integrate the original stochastic system of the SEIR model [Eqs. (1a)–(1c)] along with the transformed
stochastic system [Eqs. (20a)–(20c)] using a stochastic fourth-order Runge–Kutta scheme with a constant
time step size. The original system is solved for ,
, and ,
while the transformed system is solved for ,
, and . In
the latter case, once the values of ,
, and are
known, we compute the values of , , and
using the transformation given by
Eqs. (7a)–(7c). We shift
, , and
, respectively, by
,
,
and
to find the values of , , and
.Figure 2 compares the time series of the fraction of
the population that is infected with a disease , computed using the
original stochastic system of equations of the SEIR model with the time series of
computed using the transformed
stochastic system of equations of the SEIR model.
FIG. 2.
Time series of the fraction of the population that is infected with a disease
, computed using the original
stochastic system of equations of the SEIR model [Eqs. (1a)–(1c)] (red solid line), and computed using the transformed
stochastic system of equations of the SEIR model [Eqs. (20a)–(20c)] (blue dashed line). The standard deviation of
the noise intensity used in the simulation is ,
.
Although the two time series shown in Fig. 2
generally agree very well, there is some discrepancy. This discrepancy is due to the fact
that the noise processes ,
,
and
of the transformed system are new, independent noise processes with different variance
than the ,
,
and
noise processes found in the original system. If we carefully take the noise realization
from the original system, transform this noise using Eqs. (21a)–(21c), and use this realization to solve the
transformed system, then the two solutions would be identical.
Reduction in the stochastic SEIR model
It is tempting to consider the reduced stochastic model found by substitution of Eq.
(17) into Eqs. (20b) and (20c), so that one has the
following stochastic evolution equations (that hopefully describe the dynamics on the
stochastic center manifold):One should note that Eqs. (22a) and
(22b) also can be found by naïvely adding the stochastic terms to the reduced
system of evolution equations for the deterministic problem [Eqs. (18a) and (18b)]. This type of stochastic
center manifold reduction has been done for the case of discrete noise. Additionally, in many other fields (e.g.,
oceanography, solid mechanics, fluid mechanics), researchers have performed stochastic
model reduction using a Karhunen–Loève expansion (principal component analysis, proper
orthogonal decomposition).
However, this linear projection does not properly capture the nonlinear effects.
Furthermore, one must subjectively choose the number of modes needed for the expansion.
Therefore, even though the solution to the reduced model found using this technique may
have the correct statistics, individual solution realizations will not agree with the
original, complete solution.We will show that Eqs. (22a) and
(22b) do not contain the correct projection of the noise onto the center
manifold. Therefore, when solving the reduced system, one does not obtain the correct
solution. Such errors in stochastic epidemic modeling impact the prediction of disease
outbreak when modeling the spread of a disease in a population.Using the same numerical scheme previously described, we numerically integrate the
complete stochastic system of transformed equations of the SEIR model [Eqs. (20a)–(20c)] along with the reduced
system of equations that is based on the deterministic center manifold with a replacement
of the noise terms [Eqs. (22a) and
(22b)]. The complete system is solved for ,
, and ,
while the reduced system is solved for and
. In the latter case,
is computed using the center manifold
equation given by Eq. (17). Once the values
of , , and
are known, we compute the values of
, , and
using the transformation given by
Eqs. (7a)–(7c). We shift
, , and
, respectively, by
,
,
and
to find the values of , , and
.Figures 3(a) and 3(b) compare the time series
of the fraction of the population that is infected with a disease
, computed using the complete
stochastic system of transformed equations of the SEIR model [Eqs. (20a)–(20c)] with the time series of
computed using the reduced system of
equations of the SEIR model that is based on the deterministic center manifold with a
replacement of the noise terms [Eqs. (22a)
and (22b)]. Figure 3(a) shows the initial
transients, while Fig. 3(b) shows the time series
after the transients have decayed. One can see that the solution computed using the
reduced system quickly becomes out of phase with the solution of the complete system. Use
of this reduced system would lead to an incorrect prediction of the initial disease
outbreak. Additionally, the predicted amplitude of the initial outbreak would be
incorrect. The poor agreement, both in phase and amplitude, between the two solutions
continues for long periods of time, as seen in Fig. 3(b). We also have computed the cross correlation of the two time series shown
in Figs. 3(a) and 3(b) to be approximately 0.34.
Since the cross correlation measures the similarity between the two time series, this low
value quantitatively suggests a poor agreement between the two solutions.
FIG. 3.
Time series of the fraction of the population that is infected with a disease
, computed using the complete
stochastic system of transformed equations of the SEIR model [Eqs. (20a)–(20c)] (red solid line), and
computed using the reduced system of equations of the SEIR model that is based on the
deterministic center manifold with a replacement of the noise terms [Eqs. (22a) and (22b)] (blue dashed line). The
standard deviation of the noise intensity used in the simulation is
,
.
The time series is shown for (a)
to
and for (b)
to .
Using the same systems of transformed equations, we compute 140 years worth of time
series for 500 realizations. Ignoring the first 40 years of transient solution, the data
are used to create a histogram representing the probability density
of the and
values. Figure 4(a) shows the histogram associated
with the complete stochastic system of transformed equations, while Fig. 4(b) shows the histogram associated with the reduced
system of equations with a replacement of the noise terms. The color-bar values in Figs.
4(a) and 4(b) have been normalized by
.
FIG. 4.
Histogram of probability density
of the and
values found using (a) the complete stochastic system of transformed equations for the
SEIR model [Eqs. (20a)–(20c)]
and (b) the reduced system of equations of the SEIR model that is based on the
deterministic center manifold with a replacement of the noise terms [Eqs. (22a) and (22b)]. The histograms are
created using 100 years worth of time series (starting with year 40) for 500 realizations,
and the color-bar values have been normalized by .
One can see by comparing Fig. 4(a) with Fig. 4(b) that the two probability distributions
qualitatively look the same. It is also possible to compare the two distributions using a
quantitative measure. The Kullback–Leibler divergence or relative entropy measures the
difference between the two probability distributions aswhere
refers to the
component of the probability density found using the complete stochastic system of
transformed equations [Fig. 4(a)] and
refers to the
component of the probability density found using the reduced system of equations [Fig.
4(b)]. In our numerical computation of the
relative entropy, we have added
to each
and .
This eliminates the possibility of having a
in the denominator of Eq. (23) and does not
have much of an effect on the relative entropy.If the two histograms were identical, then the relative entropy given by Eq. (23) would be .
The two histograms shown in Figs. 4(a) and 4(b)
have a relative entropy of ,
which means that the two histograms, while not identical, are quantitatively very similar.
However, one cannot rely entirely on the histograms alone to say that the solutions of the
complete system and the reduced system agree. As we have seen in Figs. 3(a) and 3(b),
the two solutions have differing amplitudes and are out of phase with one another. It is important to note that these
features are not picked up by the histograms of Fig. 4.
CORRECT PROJECTION OF THE NOISE ONTO THE STOCHASTIC CENTER MANIFOLD
To project the noise correctly onto the center manifold, we will derive a normal form
coordinate transform for the complete stochastic system of transformed equations of the SEIR
model given by Eqs. (20a)–(20c).
The particular method we use to construct the normal form coordinate transform not only
reduces the dimension of the dynamics, but also separates all of the fast processes from all
of the slow processes. This technique
has been modified and applied to the large fluctuations of multiscale problems.Many publications
discuss the simplification of a stochastic dynamical system using a stochastic normal form
transformation. In some of this work, anticipative noise processes appeared in the normal form
transformations, but these integrals of the noise process into the future were not dealt
with rigorously.Later, the rigorous theoretical analysis needed to support normal form coordinate
transforms was developed in Refs. 23 and 24. The
technical problem of the anticipative noise integrals also was dealt with rigorously in this
work. Even later, another stochastic normal form transformation was developed. This new method allows for the “[removal
of] anticipation… from the slow modes with the result that no anticipation is required after
the fast transients decay” (Ref. 25, p. 13). The
removal of anticipation leads to a simplification of the normal form. Nonetheless, this
simpler normal form retains its accuracy with the original stochastic system.We shall use the method of Ref. 25 to simplify our
stochastic dynamical system to one that emulates the long-term dynamics of the original
system. The method involves five principles, which we recapitulate here for
completeness:Avoid unbounded
secular terms in both the transformation and the evolution equations to ensure a
uniform asymptotic
approximation.Decouple all of the slow
processes from the fast processes to ensure a valid long-term
model.Insist that the stochastic slow
manifold is precisely the transformed fast processes coordinate being equal to
zero.To simplify matters, eliminate as
many as possible of the terms in the evolution
equations.Try to remove all fast
processes from the slow processes by avoiding as much as possible the fast time memory
integrals in the evolution equations.In practice, the original stochastic system of equations (which satisfies the necessary
spectral requirements) in
coordinates is transformed to a new
coordinate system using a near-identity stochastic coordinate transform given
aswhere
the specific form of ,
,
and
is chosen to simplify the original system according to the five principles listed
previously, and is found using an iterative procedure. To outline the procedure, we provide
details for a simple example in Appendix A.Several iterations lead to coordinate transforms for ,
, and along
with evolution equations describing the -dynamics,
-dynamics,
and -dynamics
in the new coordinate system. The -dynamics have
exponential decay to the
slow manifold. Substitution of
leads to the coordinate transformswhereandAll of the stochastic terms in Eqs. (25a)–(25c) consist of integrals of the noise process into the past
(convolutions), as given by Eqs. (26) and
(27). These memory integrals are fast-time processes. Since we are interested in
the long term slow processes and since the expectation of equals ,
where ,
we neglect the memory integrals and the higher-order multiplicative terms found in Eqs.
(25a)–(25c) so
thatNote
that Eq. (28a) is the deterministic center
manifold equation, and at first order, matches the center manifold equation that was found
previously [Eq. (17)].Substitution of
and neglecting all multiplicative noise terms and memory integrals using the argument from
above (so that we consider only first-order noise terms) leads to the following reduced
system of evolution equations on the center manifold:The
specific form of and in
Eqs. (29a) and (29b) is complicated and
is therefore presented in Appendix B.We numerically integrate the complete stochastic system of transformed equations of the
SEIR model [Eqs. (20a)–(20c)]
along with the reduced system of equations that is found using the stochastic normal form
coordinate trans form [Eqs. (29a),
(29b), (B1a), and (B1b)]. The complete system is solved for
, , and
, while the reduced system is solved for
and .
In the latter case, is computed using the center manifold
equation given by Eq. (28a). As before, once
the values of , , and
are known, we compute the values of
, , and
using the transformation given by Eqs.
(7a)–(7c). We shift
, , and
, respectively, by
,
,
and
to find the values of , , and
.Figures 5(a) and 5(b) compare the time series of
the fraction of the population that is infected with a disease ,
computed using the complete stochastic system of transformed equations of the SEIR model
[Eqs. (20a)–(20c)] with the time
series of computed using the reduced system of
equations of the SEIR model that is found using the stochastic normal form coordinate
transform [Eqs. (29a), (29b), (B1a),
and (B1b) ]. Figure 5(a) shows the initial
transients, while Fig. 5(b) shows the time series
after the transients have decayed. One can see that there is an excellent agreement between
the two solutions. The initial outbreak is successfully captured by the reduced system.
Furthermore, Fig. 5(b) shows that the reduced system
accurately predicts recurrent outbreaks for a time scale that is orders of magnitude longer
than the relaxation time. This is not surprising since the solution decays exponentially
throughout the transient and then remains close to the lower-dimensional center manifold.
Since we are not looking at periodic orbits, there are no secular terms in the asymptotic
expansion, and the result is valid for all time. Additionally, any noise drift on the center
manifold results in bounded solutions due to sufficient dissipation transverse to the
manifold. The cross correlation of the two time series shown in Fig. 5 is approximately 0.98, which quantitatively suggests there is an
excellent agreement between the two solutions.
FIG. 5.
Time series of the fraction of the population that is infected with a disease
, computed using the complete
stochastic system of transformed equations of the SEIR model [Eqs. (20a)–(20c)] (red solid line), and
computed using the reduced system of equations of the SEIR model that is found using the
stochastic normal form coordinate transform [Eqs. (29a), (29b), (B1a), and (B1b)] (blue dashed line). The
standard deviation of the noise intensity used in the simulation is
,
.
The time series is shown for (a)
to
and for (b)
to .
Using the same systems of transformed equations, we compute 140 years worth of time series
for 500 realizations. As before, we ignore the first 40 years worth of transient solution,
and the data are used to create a histogram representing the probability density
of the and
values. Figure 6(a) shows the histogram associated
with the complete stochastic system of transformed equations, while Fig. 6(b) shows the histogram associated with the reduced
system of equations found using the normal form coordinate transform. The color-bar values
in Figs. 6(a) and 6(b) have been normalized by
.
FIG. 6.
Histogram of probability density
of the and
values found using (a) the complete stochastic system of transformed equations for the
SEIR model with mortality [Eqs. (20a)–(20c)] and (b) the reduced system of equations of the SEIR model with
mortality that is found using the stochastic normal form coordinate transform [Eqs. (29a), (29b), (B1a), and
(B1b)]. The histograms are created using 100 years worth of time series (starting
with year 40) for 500 realizations, and the color-bar values have been normalized by
.
As we saw with Figs. 4(a) and 4(b), the
probability distribution shown in Fig. 6(a) looks
qualitatively the same as the probability distribution shown in Fig. 6(b). Using the Kullback–Leibler divergence given by Eq. (23), we have found that the two histograms shown
in Figs. 6(a) and 6(b) have a relative entropy of
.
Since this value is close to zero, the two histograms are quantitatively very similar.In addition to computing the cross correlation between the solution of the original system
and the solutions of the two reduced systems for ,
we have computed the cross correlation for the case of zero noise as well as for noise
intensities ranging from
to .
These cross correlations were computed using time series from
to .
For the deterministic case (zero noise), the cross correlation between the time series which
were computed using the original system and the reduced system based on the deterministic
center manifold is 1.0, since the agreement is perfect. The cross correlation between the
original system and the reduced system found using the stochastic normal form is also 1.0.
Figure 7 shows the cross correlation between the
original system and the two reduced systems for various values of .
FIG. 7.
Cross correlation between time series found using the original stochastic system of
transformed equations and the reduced system of equations based on the deterministic
center manifold (“circle” markers) and cross correlation between time series found using
the original stochastic system of transformed equations and the reduced system of
equations based on the stochastic normal form coordinate transform (“square” markers). The
cross correlation is computed using time series from
to .
One can see in Fig. 7 that the solutions found using
the reduced system based on the deterministic center manifold compare poorly with the
original system at very low noise values. Furthermore, as the noise increases, the agreement
between the two solutions gets worse. On the other hand, Fig. 7 shows that the solutions computed using the reduced system found using the
normal form coordinate transform compare very well with the solutions to the original system
across a wide range of small noise intensities.
DISCUSSION
We have demonstrated that the normal form coordinate transform method reduces the Langevin
system so that both the noise and dynamics are accurately projected onto the
lower-dimensional manifold. It is natural to consider (a) the replacement of the stochastic
term by a deterministic period drive of small amplitude and (b) the extension to finite
populations. These cases are discussed, respectively, in Secs. VI A and VI B.
The case of deterministic forcing
A single time series realization of the noise might be thought of as a deterministic
function of small amplitude driving the system. One could rederive the normal form for
such a deterministic function. However, since our derived normal form holds specifically
for the case of white noise, we show that a simple replacement of the stochastic
realization with a deterministic realization does not work. As an example, one could
consider the following sinusoidal functions:where
,
,
and
are given by Eqs. (21a)–(21c).
Using Eqs. (30a)–(30c) or some
other similar deterministic drive, the solution computed using the reduced system based on
the deterministic center manifold analysis will agree perfectly with the solution computed
using the complete system of equations. On the other hand, since the reduced system based
on the normal form analysis was derived specifically for white noise, the transient
solution found using this reduced system will not agree with the solution found using the
complete system. It is possible to find a normal form coordinate transform for periodic
forcing, but the normal form will be different than the one derived in this article for
white noise.Figures 8(a) and 8(b) compare the time series
of the fraction of the population that is infected with a disease
, computed using the complete system of
transformed equations of the SEIR model [Eqs. (20a)–(20c)] with the time series of
computed using the reduced system of
equations of the SEIR model that is found using the stochastic normal form coordinate
transform [Eqs. (29a), (29b),
(B1a), and (B1b)], but where the stochastic terms of both systems have been
replaced by the deterministic terms given by Eqs. (30a)–(30c). Figure 8(a) shows the initial transients, while Fig. 8(b) shows a piece of the time series after the transients have decayed. One can
see in Figs. 8(a) and 8(b) that although the two
solutions eventually become relatively synchronized with one another, there is a poor
agreement, both in phase and amplitude, throughout the transient.
FIG. 8.
Time series of the fraction of the population that is infected with a disease
, computed using the complete system of
transformed equations of the SEIR model [Eqs. (20a)–(20c)] (red solid line), and computed using the
reduced system of equations of the SEIR model that is found using the normal form
coordinate transform [Eqs. (29a),
(29b), (B1a), and (B1b)] (blue dashed line). The stochastic terms in both systems
have been replaced by the deterministic terms given by Eqs. (30a) and (30b). The time series is shown from (a)
to
and from (b)
to .
The case of finite populations
The solutions to the original system and both reduced systems are continuous solutions
based on an infinite population assumption and are found using Langevin equations having
Gaussian noise. It is interesting to examine the effects of general noise by using a
Markov simulation to compare solutions of the original and reduced systems.The complete system in the original variables (see p. 2) will evolve in time
in the following way:Using
a total population size of ,
we have performed a Markov simulation of the system. After completing the Markov
simulation, we divided , , and
by to
find , , and
. Figure 9(a) shows a time series, after the transients have decayed, of the fraction of
the population that is infected with a disease . The results reflect
both the mean and the frequency of the deterministic system. Performing the simulation for
500 realizations allows us to create a histogram representing the probability density
of the and
values. This histogram is shown in Fig. 9(b), and
one can see that the probability density reflects the amplitude, which varies with the
population size of and . The
color-bar values in Fig. 9(b) have been normalized
by
FIG. 9.
(a) Time series of the fraction of the population that is infected with a disease
, computed using a Markov simulation of
the complete original equations of the SEIR model [Eq. (31)] and (b) a histogram of probability density
of the and
values found using a Markov simulation of Eq. (31). The histogram is created using 100 years worth of data (starting with year
40) for 500 realizations, and the color-bar values have been normalized by
.
The complete system in the transformed variables has the stable endemic equilibrium at
the origin. To bound the dynamics to the first octant, we use the fact that
,
,
and
to derive the appropriate inequalities for the transformed discrete variables
, , and
. These inequalities can be found in
Appendix C as Eq. (C1). These inequalities enable us to define new discrete variables
,
,
and
given by Eqs. (C2a)–(C2c) in Appendix C.In the
variables, we define evolution relationships similar to those found in Eq. (31). The complete transformed system will
evolve in time according to the transition and
rates given by Eq. (C3) in Appendix C.After performing a Markov simulation of Eq. (C3) with a population size of ,
we can compare the dynamics of the transformed system with the dynamics of the original
system by transforming the
variables in the time series back to the original ,
, and
variables. Dividing by yields ,
, and .
Figure 10(a) shows a time series, after the
transients have decayed, of the fraction of the population that is infected with a disease
. The mean and the frequency agree with
those found from the Markov simulation of the original system. We have performed the
simulation for 500 realizations, and a histogram representing the probability density
is shown in Fig. 10(b). The color-bar values in
Fig. 10(b) have been normalized by
.
One can see in Fig. 10(a) that the relative
fluctuations of the component have nearly doubled. While
the fluctuation size was 0.152 for the original system, it is 0.310 for the transformed
system. Additionally, the two histograms shown in Figs. 9(b) and 10(b) have a
relative entropy of ,
which means they are not in agreement. Because the simulation of the stochastic dynamics
in the complete system of transformed variables does not qualitatively (or quantitatively)
resemble the original stochastic system, we cannot expect that the reduced system will
agree with either the original or the transformed systems. Therefore, much care should be
exercised when extending the model reduction results (which show outstanding agreement)
derived for a specific type of noise in the limit of infinite population to finite
populations with a more general type of noise.
FIG. 10.
(a) Time series of the fraction of the population that is infected with a disease
, computed using a Markov simulation of
the complete transformed equations of the SEIR model [Eq. (C3)] and (b) a histogram of probability density
of the and
values found using a Markov simulation of Eq. (C3). The histogram is created using 100 years worth of data (starting with year
40) for 500 realizations, and the color-bar values have been normalized by
.
CONCLUSIONS
We have considered the dynamics of a SEIR epidemiological model with stochastic forcing in
the form of additive Gaussian noise. We have presented two methods of model reduction,
whereby the goal is to project both the noise and the dynamics onto the stochastic center
manifold. The first method uses the deterministic center manifold found by neglecting the
stochastic terms in the governing equations, while the second method uses a stochastic
normal form coordinate transform.Since the original system of governing equations does not have the necessary spectral
structure to employ either deterministic or stochastic center manifold theory, the system of
equations has been transformed using an appropriate linear transformation coupled with
appropriate parameter scaling. At this stage, the first method of model reduction can be
performed by computing the deterministic center manifold equation. Substitution of this
equation into the complete stochastic system of transformed equations leads to a reduced
system of stochastic evolution equations.The solutions of the complete stochastic system of transformed equations as well as the
reduced system of equations were computed numerically. We have shown that the individual
time series does not agree because the noise has not been correctly projected onto the
stochastic center manifold. However, by comparing histograms of the probability density
of the and
values, we saw that there was a very good agreement. This is caused by the fact that
although the two solutions are out of phase with one another, their range of amplitude
values is similar. The phase difference is not represented in the two histograms. This is a
real drawback when trying to predict the timing of outbreaks and leads to potential problems
when considering epidemic control, such as the enhancement of disease extinction through
random vaccine control.To accurately project the noise onto the manifold, we derived a stochastic normal form
coordinate transform for the complete stochastic system of transformed equations. The
numerical solution to this reduced system was compared with the solution to the original
system, and we showed that there was an excellent agreement both qualitatively and
quantitatively. As with the first method, the histograms of the probability density
of the and values
agree very well.It should be noted that the use of these two reduction methods is not constrained to
problems in epidemiology, but rather may be used for many types of physical problems. For
some generic systems, such as the singularly perturbed, damped Duffing oscillator, either
reduction method can be used since the terms in the normal form coordinate transform which
lead to the average stochastic center manifold being different from the deterministic center
manifold occur at very high order. In
other words, the average stochastic center manifold and deterministic center manifold are
virtually identical. For the SEIR model considered in this article, there are terms at low
order in the normal form transform, which cause a significant difference between the average
stochastic center manifold and the deterministic manifold. Therefore, as we have
demonstrated, when working with the SEIR model, one must use the normal form coordinate
transform method to correctly project the noise onto the center manifold.In summary, we have presented a new method of stochastic model reduction that allows for
impressive improvement in time series prediction. The reduced model captures both the
amplitude and phase accurately for a temporal scale that is many orders of magnitude longer
than the typical relaxation time. Since sufficient statistics of disease data are limited
due to short time series collection, the results presented here provide a potential method
to properly model real, stochastic disease data in the time domain. Such long-term accuracy
of the reduced model will allow for the application of effective control of a disease where
phase differences between outbreak times and vaccine controls are important. Additionally,
since our method is general, it may be applied to very high-dimensional epidemic models,
such as those involving adaptive networks. From a dynamical systems viewpoint, the reduction
method has the potential to accurately capture new, emergent dynamics as we increase the
size of the random fluctuations. This could be a means to identify new noise-induced
phenomena in generic stochastic systems.