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ISSN 1979-3898

Journal of Theoretical and Computational Studies

**Simple Dynamics in a Vector-Borne Disease Model
**

A.K. Supriatna and E. Soewono J. Theor. Comput. Stud. 8 (2008) 0401 Received: July 7th , 2008; Accepted for publication: September 8th , 2008

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**

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J. Theor. Comput. Stud. Volume 8 (2008) 0401

**Simple Dynamics in a Vector-Borne Disease Model
**

Asep K. Supriatna Department of Mathematics, Universitas Padjadjaran, Jl. Raya Bandung-Sumedang Km 21, Jatinangor 45363, Indonesia Edy Soewono Industrial and Financial Mathematics Group, Institut Teknologi Bandung, Jl. Ganesha 10, Bandung 40132, Indonesia

Abstract : In this paper we review a simple model of an infectious disease transmission. In general the rate of incidences can be model by mass action principle, so that its functional is bilinear. In some circumstances, disease transmission might be more complicated involving diﬀerent species, for example in the case of the transmission of the disease required a vector (vector-borne disease), such as in malaria and dengue infection cases, the rate of incidences takes a nonlinear functional form. In this paper we show the conditions needed for the endemic equilibrium in the model to exist and to be stable. The analysis reveals that there is a simple transcritical bifurcation in the dynamics of the model, despite the complex interaction of the disease transmission. Keywords : Epidemic model, host-vector transmission, basic reproduction number, population dynamics E-mail : aksupriatna@bdg.centrin.net.id Received: July 7th , 2008; Accepted for publication: September 8th , 2008

1

INTRODUCTION

Despite recognized as an abstract science, mathematics has proved to be useful in helping to solve many problems arising in daily life and problems from other disciplines, such as industrial, environmental, and biological sciences (see [17, 16] and the reference therein for examples). The inter-relations between mathematics and other disciplines not merely have given beneﬁts to the disciplines served by mathematics, but in many cases, there also fruitfulness to mathematics itself. There are some mathematical concepts and theories inspired from these inter-relations. Sometimes the intimate connection between mathematics and other discipline gives rise to a new discipline, such as mathematical bio-economics [5], mathematical conservation theory [4] and mathematical epidemiology [1]. In this paper we will review an application of mathematics in controlling the transmission of an infectious disease. The ﬁrst documented work on the application of mathematics in controlling an infectious disease dates back to the 18th century when Daniel Bernoulli used a mathematical method to evaluate the eﬀectiveness of the techniques of variolation against smallpox [2]. Among the aims of his work was to inﬂuence the pubc 2008 GFTI & MKI

lic health authority in deciding the eﬀectiveness of the infectious disease control at the time. He showed that the techniques of inoculation practiced by the society could increase the number of survivors per year or increase the average life expectancy in an epidemic episode, if it is implemented universally to the whole population (known as a universal inoculation method). Current review on his work can be found in [9]. Early scientists postulated that the course of an epidemic depends on the rate of contact between susceptible and infected individuals. Generally, they model this phenomenon through the mass action principle. Among them is Ross [15], a medical doctor who served as a colonel in the British Army. He used this principle, identiﬁed the main factors in malaria transmission, calculated the number of new infection, and concluded that no need to eradicate all of the mosquitoes to eradicate malaria, because there exists a critical density of mosquito, below which the disease will vanish. This result is usually known as the mosquito theorem or the theory of critical level density. His work is then generalized extensively by Kermack and McKendrick [11]. Current review on their work can be found in [8, 10, 3]. 0401-1

2 A concept similar to the theory of critical level density is introduced by McDonald [14] who coined it as the basic reproduction rate. In a modern notation this concept is symbolized by R0 and is deﬁned as the expected number of secondary cases produced, in a completely susceptible population, by a typical infected individual during its entire period of infectiousness [7, 6]. In the following sections we will discuss a simple epidemic model and show that in some circumstances nonlinearity may often occur as a result of a complex epidemiological phenomenon. 2 2.1 SIMPLE EPIDEMIC MODELS A model without demographic parameter

Simple Dynamics in a Vector-Borne... eventually the disease will die out leaving a portion of uninfected population, regardless the value of the initial conditions (see Figure 1 for illustration). The general solution of the system can also be found dS in the SR-plane by looking at the equation dR = −βIS S (−R/ρ) as the solution γI = − ρ which gives S = S0 e through (S0 , I0 ). If the initial infection is relatively low, I0 ≈ 0 then considering the steady state solution of (1)-(3), we have R ≈ 2ρ(1 − ρ/S0 ). Furthermore, if ε S0 = ρ + ε with ε → 0 then R ≈ 2ρ ρ+ε ≈ 2ε. Considering the intensity of the epidemic is measured by R(∞) the expression R ≈ 2ε means that the epidemic has succeeded in reducing the density of susceptible from the initial condition ρ + ε to the ﬁnal condition ρ − ε. This is known as the Kermack-McKendrick threshold theorem. 2.2 A model with demographic parameters

A fairly simple epidemic model is an SIR model in which we divide the population into three compartments, namely the susceptible (S), infected (I) and recover (R) sub-populations. Here we assume that the total population (N ) is constant with N = S + I + R. If it is further assumed that the force of infection is β and removal rate is γ then the dynamics of the disease transmission is given by dS = −βIS, dt (1)

To increase realism, demographic parameters, such as birth rate and death rate, are incorporated into the model in the previous section. Suppose that the birth rate is a constant B, and the death rate is proportional to the population density, with the constant of proportionality µ. Hence, the system becomes dS = B − βIS − µS, dt (4)

this regard, R0 = β S0 is called the basic reproduction γ number of the system. If the value of this basic reproduction number is more than one, then the number of infected population increases, otherwise the number of infected population decreases. The general solution of the system can be found dI in the SI-plane by looking at the equation dS = βIS−γI = −1 + ρ/S. Next, by considering N0 = −βIS S0 + I0 , the solution through (S0 , I0 ) is given by S I = N − S + ρ ln S0 , having its maximum infection Imax = N − ρ + ρ ln(ρ/S0 ) at S = ρ. Furthermore, it can be shown that I(∞) = 0 and S(∞) > 0 found as the smallest positive root of equation N − S(∞) + ρ log [S (∞)/S0 ] = 0. This means that

dI = βIS − γI, (2) dt dR = γI, (3) dt with initial conditions S(0) = S0 > 0, I(0) = I0 > 0, and R(0) = 0. Note from (2) that in the beginning of the epidemic, there will be a build up of infection rises to an outbreak only if S0 > ρ = γ/β, otherwise the epidemic will die out. Hence, S0 = ρ is a threshold density of susceptible. Here γ/β is the relative removal rate. This threshold can be reformulated as follows. The condition S0 > ρ = γ/β is equivalent to β S0 > 1. In γ

dI = βIS − γI − µI, (5) dt dR = γI − µR. (6) dt Now, from (5), there will be a build up of infection rises to an outbreak only if S0 > ρµ = (γ + µ)/β, otherwise the epidemic will die out. In either case, there are two equilibrium solutions possible ∗ ∗ to occur, endemic-free equilibrium state (S0 , I0 ) = B ∗ ∗ and endemic equilibrium state (S1 , I1 ) = µ,0

γ+µ 1 β , β B ∗ S1

−µ

. However, the endemic equilib-

B rium state only appears when S ∗ − µ > 0 or equiv1 alently its basic reproduction number is more than Bβ one, that is R0 = µ(γ+µ) > 1. This basic reproduction number is also a threshold for stability, in the sense that the endemic equilibrium state exists and is stable if R0 > 1 while the endemic-free equilibrium is unstable, otherwise the endemic equilibrium state does not exist while the endemic-free equilibrium state is stable. Note that the condition of R0 > 1 is a suﬃcient condition for the endemic equilibrium to occur, while the inclusion demographic parameter B is a necessary condition. Meanwhile, the inclusion of demographic

0401-2

3

Simple Dynamics in a Vector-Borne...

1

1

In(t)

0,8

In(t)

0,8

0,6

0,6

0,4

0,4

0,2

0,2

0 0 0,2 0,4 0,6 0,8 S(t) 1

0 0 0,2 0,4 0,6 0,8 S(t) 1

Figure 1: The trajectory of the SIR system without demographic parameter in the SI-plane. It shows that eventually the disease will die out regardless the value of the initial conditions. The parameter values in the ﬁgure are γ = 0.3 and β = 1 with various initial values of S and I.

parameter µ changes the value of the threshold density of susceptible. An illustration of the solutions for various initial conditions can be seen in Figure 2. 3 A MODEL FOR VECTOR-BORNE DISEASE TRANSMISSION

Figure 2: The trajectory of the SIR system in the SIplane with the inclusion of demographic parameters and R0 > 1. It shows that the disease will be endemic eventually. The parameter values in the ﬁgure are the same as in ﬁgure 1 with additions B = 0.08 and µ = 0.1. The resulting basic reproduction number is R0 = 2 indicating the endemicity of the disease.

dSV = BV − βV IH SV − µV SV , dt dIV = βV IH SV − µV IV , dt

(10) (11)

Many diseases required a vector to spread. For example mosquitoes are responsible in dengue and malaria transmission. In regards to the transmission of a vector-borne disease, the previous models ignore the presence of vectors in spreading the disease. In this section we generalize the previous model with demographic parameters to include a vector in the model. Let us assume SH (t) be the density of susceptible human population, IH (t) be the density of infected human population, RH (t) be the density of removed and immune human population, SV (t) be the density of susceptible vector population, and IV (t) be the density of infected vector population. The governing equations for the transmission of the disease in the presence of demographic parameters are dSH = B − βH IV SH − µH SH , dt dIH = βH IV SH − IH (γH + µH ) , dt dRH = γH IH − µH RH , dt (7) (8) (9)

B ∗ IV = µV (βV βV IH V ) . Substituting these values to the V IH +µ human dynamics yields approximation equations to the original vector-borne disease transmission

where, as before, we also assume SH + IH + RH = NH and SV + IV = NV . Following [12], we further assume that the vector dynamics runs on a much faster time scale than the human dynamics. Hence the vector population can be considered to be at its equilibrium with the respect to the changes in human population. Equilibrium solu∗ tions for the vector is given by SV = βV IBV V and H +µ

βH BV βV IH SH dSH =B− − µH SH , dt µV (βV IH + µV ) dIH βH BV βV IH SH = − IH (γH + µH ) , dt µV (βV IH + µV ) dRH = γH IH − µH RH . dt

(12)

(13) (14)

Compared to the direct transmission model, in which the incidence rate is a bilinear function, here the inci-

0401-3

4

Simple Dynamics in a Vector-Borne...

1

1

In(t)

0,8

In(t)

0,8

0,6

0,6

0,4

0,4

0,2

0,2

0 0 0,2 0,4 0,6 0,8 S(t) 1

0 0 0,2 0,4 0,6 0,8 S(t) 1

Figure 3: The trajectory of the SIR system for the hostvector model in the SI-plane with various initial values of S and I. The parameter values in the ﬁgure are γH = 0.3, βH = 1, BH = 0.08, µH = 0.1, βV = 1, BV = 0.1, and µV = 0.5. The basic reproduction number is R0 = 0.8 indicating the vanishing of the disease.

Figure 4: The trajectory of the SIR system for the hostvector model in the SI-plane with various initial values of S and I, with the same parameters as in ﬁgure 3 except a lower vector mortality rate, i.e. µV = 0.1. The resulting basic reproduction number is R0 = 20 indicating the endemicity of the disease.

dence rate f (IH , SH ) = βH BV βV IH SH µV (βV IH + µV ) (15)

∗ ∗ ∗ is nonlinear in IH . It is clear that P0 = (SH0 , IH0 ) =

is the endemic-free equilibrium solution. The ∗ ∗ endemic equilibrium solution is given by p∗ (SHe , IHe ) e in which

∗ SHe

B µH , 0

Next, following [13], we derive lemmas relating the basic reproduction number R0 with the properties of the nonlinear incidence rate f (SH , IH ) at the equlibria of (12)-(14). Lemma 3.1: At the endemic-free equilibrium state ∗ ∗ p∗ = (SH0 , IH0 ), we have 0 R0 ≤ 1 ⇒ R0 > 1 ⇒

∗ ∗ ∂f (SH0 , IH0 ) ≤ γH + µ H , ∂IH ∗ ∗ ∂f (SH0 , IH0 ) > γH + µH . ∂IH

BβV µV + µ2 (γH + µH ) H = , (BV βV βH + µH µV βV )

∗ IHe = (R0 − 1) X ∗ ,

(16) (17)

Proof: In relation to the incidence rate in (15), the basic reproduction number in (19) can be written as R0 = = BBV βV βH µH µ2 (γH + µH ) V ∂f (IH , SH ) 1 (γH + µH ) ∂IH

with X∗ = µH µ2 (γH + µH ) V (γH + µH ) (BV βV βH + µH µV βV ) R0 = BBV βV βH . µH µ2 (γH + µH ) V , (18) (19)

.

∗ ∗ (IH ,SH )=(I0 ,S0 )

It is easy to see that the nonlinear incidence rate f (SH , IH ) = βH BV βV IH SH has the following properµV (βV IH +µV ) ties: P1: f (0, IH ) = f (SH , 0) = 0, ,I P2: ∂f (SHH H ) > 0 and ∂f (SH ,IH ) > 0, ∂S ∂IH P3:

∂ 2 f (SH ,IH ) 2 ∂IH

Hence, the proof of the lemma is clear. Lemma 3.2: At the endemic equilibrium state p∗ = e ∗ ∗ (SHe , IHe ), we have R0 > 1 ⇒ Furthermore,

∗ ∗ ∂ 2 f (SHe ,IHe ) 2 ∂IH

∗ ∗ ∂f (SHe , IHe ) ≤ γH + µH . ∂IH

∗ ∗ ∂f (SHe ,IHe ) ∂IH

=

γH + µH only if

≤ 0.

= 0.

0401-4

5 Proof: We observe, from (13), that the en∗ ∗ demic equilibrium state satisﬁes f (SHe , IHe ) = ∗ ˜(IH ) := f (S ∗ , IH ). (γH + µH ) IHe . Let us deﬁne f He∗ ∗ ∂f (SHe ,IHe ) Suppose that, in the contrary, we have = ∂IH > γH + µH . By considering P1, the mean value theorem guarantees that there is a point IH1 ∈ ∗ (0, IHe ) such that ˜ df (IH1 ) dIH = = ˜ ∗ ˜ f (IHe ) − f (0) ∗ −0 IHe ∗ (γH + µH ) IHe − 0 = γH + µH . ∗ IHe

˜ df (I ∗ ) ˜ ∗ df (IHe ) dIH

Simple Dynamics in a Vector-Borne... Stability of the endemic equilibrium state is clear from the following theorem. Theorem 2: If R0 > 1 then the endemic equilib∗ ∗ ∗ rium Pe = (SHe , IHe ) of the system (12)-(14) exists and is locally asymptotically stable. Proof: It is obvious by (17) the endemic equilibrium exists only if R0 > 1. The stability of the endemic equilibrium state is investigated by observing the Jacobian matrix M of the system (12)-(14). At this equilibrium state, we have the characteristic equation λ2 + a1 λ + a2 = 0, with a1 = −trace(M ) ∂f + µH = ∂SH a2 = = det(M ) (γH + µH ) +µH ∂f ∂SH ∂f ∂IH . ∂f ∂IH

He Moreover, since we have assumed dIH > γH + µH , then again by the mean value theorem there exist a ∗ point IH0 ∈ (IH1 , IHe ) such that

+ (γH + µH ) −

,

∂

2

∗ f (SHe , IH0 ) 2 ∂IH

= = =

˜ d f (IH0 ) 2 dIH

2 ˜ ∗ ˜(IH1 df (IHe ) − dfdIH ) dIH ∗ IHe − IH1 ˜ ∗ df (IHe ) − (γH + µH ) dIH ∗ IHe − IH1

∗ ∗ ∂f (SHe ,IHe ) ∂IH

(γH + µH ) −

> 0.

From Lemma 3.2 and P2 we conclude a1 > 0 and a2 > 0 which conﬁrms that the endemic equilibrium state is asymptotically stable. ACKNOWLEDGMENTS Financial support was provided by The Royal Netherlands Academy of Arts and Sciences (KNAW). Earlier version of the paper was presented in the Workshop of Nonlinear Phenomena, Bandung 11 December 2007.

JTCS

**This contradicts P3, and hence
**

∗ ∗ ∂f (SHe ,IHe ) = Furthermore, ∂IH ˜ ∗ ˜ df (IHe ) df (IH1 ) = dIH for all IH ∈ dIH

≤ γH +µH .

**γH + µH holds only if
**

∗ ∗ (0, IHe ) and SH = SHe .

In the following section we will investigate the stability of these equilibria in relation to the basic reproduction number R0 . 3.1 Stability of the equilibrium states

REFERENCES [1] R.M. Anderson and R.M. May, Infectious Diseases of Human: Dynamics and Control, Oxford University Press (1991). [2] S. Blower, Rev. Med. Virol., 14 (2004) 275. [3] F. Brauer, Math. Biosc. 198 (2005) 119. [4] M.A. Burgman, S. Ferson and H.R. Akcakaya, Risk Assessment in Conservation Biology, Chapman & Hall, London (1994). [5] C.W. Clark, Mathematical Bioeconomics: The optimal Management of Renewable Resources, Wiley Interscience, New Jersey (1990). [6] O. Diekmann and J.A.P Heesterbeek, Mathematical Epidemiology of Infectious Diseases: Model Building, Analysis and Interpretation, John Wiley & Son, Chichester (2000). [7] O. Diekmann, J.A.P. Heesterbeek and J.J. Metz, J. Math. Biol. 28 (1990) 365.

The stability results are summarized in the following theorems. Theorem 1:If R0 < 1 then the endemic-free equi∗ ∗ librium p∗ = (SH0 , IH0 ) of the system (12)-(14) is 0 locally asymptotically stable, otherwise it is unstable. Proof: The stability of the endemic-free equilibrium is easily identiﬁed through the investigation of the Jacobian matrix M of the system (12)-(14). At the endemic-free equilibrium, we have the characteristic equation λ2 + a1 λ + a2 = 0, with a1 = −trace(M ) = µH + (γH + µH ) (1 − R0 ) , a2 = det(M ) = µH (γH + µH ) (1 − R0 ) . This means that the endemic-free equilibrium is stable if R0 < 1 and is unstable if R0 > 1. The endemicfree equilibrium undergoes a transcritical bifurcation at R0 = 1.

0401-5

6 [8] O. Diekmann, J.A.P. Heesterbeek and J.J. Metz, The legacy of Kermack and McKendrick (1995) 95-115. [9] K. Dietz and J.A.P. Heesterbeek, Math. Biosc. 180 (2002) 1. [10] H. Inaba, Kermack and McKendrick revisited: The variable susceptibility model for infectious diseases, Technical Paper, University of Tokyo (2000) 11. [11] W.O. Kermack and A.G. McKendrick, Proc. Roy. Soc. Med. A115 (1927) 700. [12] J.C. Koella and R. Antia, Malaria J. 2 (2003) 1. [13] A. Korobeinikov and P.K. Maini, Math. Med. Biol. 22 (2005) 113. [14] G. McDonald, Trop. Dis. Bull. 49 (1952) 813. [15] R. Ross, The Prevention of Malaria, John Murray, London (1911). [16] E. Soewono and A.K. Supriatna, Paradox of vaccination predicted by a simple dengue disease model, Industrial Mathematics, M.C. Joshi, A.K. Pani and S.V. Sabnis Eds., Narosa Publ. House, New Delhi, (2006) 459. [17] A.K. Supriatna and H.P. Possingham, Bull. Math. Biol. 60 (1998) 49.

Simple Dynamics in a Vector-Borne...

Presented at Workshop on Nonlinear Phenomena 2K7, Sumedang, Indonesia, December 11th , 2007.

0401-6

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