Abstract
A sexually-transmitted disease model for two strains of pathogen in a one-sex, heterogeneously-mixing population has been studied completely by Jiang and Chai in (J Math Biol 56:373–390, 2008). In this paper, we give a analysis for a SIS STD with two competing strains, where populations are divided into three differential groups based on their susceptibility to two distinct pathogenic strains. We investigate the existence and stability of the boundary equilibria that characterizes competitive exclusion of the two competing strains; we also investigate the existence and stability of the positive coexistence equilibrium, which characterizes the possibility of coexistence of the two strains. We obtain sufficient and necessary conditions for the existence and global stability about these equilibria under some assumptions. We verify that there is a strong connection between the stability of the boundary equilibria and the existence of the coexistence equilibrium, that is, there exists a unique coexistence equilibrium if and only if the boundary equilibria both exist and have the same stability, the coexistence equilibrium is globally stable or unstable if and only if the two boundary equilibria are both unstable or both stable.
Citation: Chai C, Jiang J (2011) Competitive Exclusion and Coexistence of Pathogens in a Homosexually-Transmitted Disease Model. PLoS ONE 6(2): e16467. https://doi.org/10.1371/journal.pone.0016467
Editor: Vladimir Brusic, Dana–Farber Cancer Institute, United States of America
Received: October 30, 2010; Accepted: December 19, 2010; Published: February 15, 2011
Copyright: © 2011 Chai, Jiang. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: Supported by the Youth Foundation of Anhui University of Finance and Economics No. ACKYQ1065ZC, Chinese NSF grants 10671143 and 10531030. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Introduction
An important principle in theoretical biology is that of competitive exclusion: no two species can forever occupy the same ecological niche. Classifications on the meaning of competitive exclusion and niche have been central to theoretical ecology [1]–[4]. On the other hand, biologists and mathematical modelers have long been concerned with the evolutionary interactions that result from changing host and pathogen populations. Continuous advances in biology and behavior have brought to the forefront of research the importance of their role in disease dynamics [5]–[17]. Sexually transmitted diseases, such as gonorrhea have incredibly high incidences throughout the world, providing the necessary environment and opportunities for the evolution of new strains(see [18] and the references therein). The coexistence of gonorrhea strains has become an increasingly serious problem. Understanding the mechanisms that lead to coexistence or competitive exclusion is critical to the development of disease management strategies, as well as to our understanding of STD dynamics.
In previous papers [18], [19], they have shown that coexistence of multiple strains is not possible in a heterosexually-active homogenous population where individuals have the same mean behavior by investigating SIS STD models and establishing that such populations are unable to support multiple strains. However, using simple heterosexual mixing models, Castillo-Chaves et al. [20], [21] have shown that heterogeneity(behavioral or genetically or a combination of both) of one sex population(the female population) is enough to maintain heterogeneity and to lead possible coexistence of multiple strains. Chai [22] and Qiu [23] has given the completely classification for this model. Li et al. [24] have determined what is the minimum level of heterogeneity required to support multiple strains to coexist. They formulated and analyzed a one-sex, SIS STD model with two competing strains under the same assumptions. Furthermore, in [25], we have presented a thorough classification of dynamics for this model in terms of the first and the second so called reproductive numbers, and discussed the biological meaning of our results in the finally.
This paper focus on the dynamics of sexually transmitted pathogens in a homosexually active population, where populations are divided into three groups based on their susceptibility to infection(colonization) by two distinct pathogenic strains of an STD. It is assumed that a host cannot be invaded simultaneously by both disease agents(that is, there is no superinfection) and that when symptoms appear-a function of pathogen, strain, virulence, and an individual's degree of susceptibility-then individuals are treated and/or recover.
Methods
Let, denote the susceptibles with sexual activity
, which is the number of contacts per individual in group
per unit of time, and use
and
to denote the infectives with sexual activity
and infected by strain 1 and strain 2, respectively. The dynamics of the disease transmission then is described by the following equations:
(1)where
are the rates of incidence with
being the population size of group
,
are the constant input flows entering the sexually active sub-populations,
are the average sexual life spans for people in group
,
and
are the transmission probabilities per contact with individuals infected by strains 1 and 2, respectively, and
and
are the rates of recovery for classes
and
, respectively. It is assumed that people with different sexual activity having different rates of recovery as highly sexually-active individuals may have health examinations more frequently.
The limiting system of (1) is(2)where
SetThen
With these notations, the system (2) can be rewritten into the following compact form:
(3)
Note that is the total population of group
. Throughout this paper will consider only the dynamics of (3) in
, where
and
. Let
denote the solution flow generated by (3). It is not difficult to see that the flow is positively invariant in
.
For two vectors , define the vector order as follows:
and also define the type-K order in
in the sense that
The Jacobian-matrix at each point
has the form
(4)It follows from Smith [26] that the flow
is type-K monotone in the sense that
Discussion
Next, we consider the necessary thresholds and the stability of the infection-free state, established the principle of competitive exclusion and coexistence for SIS models with heterogeneous mixing.
Thresholds
The linearization about the infection-free equilibrium of (3) iswhere
Now we define the reproductive numbers
(5)Hence, by calculation, it follows from M-matrix theory [27], if
and
, then the origin is locally asymptotically stable. If
or
, the infection-free equilibrium is unstable.
As in [24], it can be shown that the locally stable infection-free equilibrium and the locally stable boundary equilibrium associated with model (3), which will be studied in the following section, are globally stable. We only state the results as follows and omit the details. The interested reader is referred to [24].
Lemma 1.
Let
and
be equilibria of (3), where
, if
and
;
, if
and
,
. Let
and
. Then
In summary, we state the threshold conditions for the disease as follows.
Theorem 1.
Let the reproductive number and
be defined in (5). Then, if
and
, the infection-free equilibrium is globally asymptotically stable so that the epidemic goes extinct regardless of the initial levels of infection. If
or
, then the infection-free equilibrium is unstable and the epidemic spreads in the population.
The computation of boundary equilibria
Let and
. Then
are invariant for (3). The subsystems on
and
are
(3)_{\it I})and
(3)_{\it J})respectively.
Following Smith [28], both and
are strongly concave. From [28] it follows that the origin is globally asymptotically stable, or there is exists and equilibrium
with
such that it is globally asymptotically stable in
. Moreover,
is also linearly stable, that is,
has the following form
is stable matrix.
From Theorem 1, if , then the origin is globally asymptotically stable in
, otherwise,
,
exists. Next, we discuss the computation for
for the case
. Make the transformation
(6)where
Then
satisfy the equations
(7)where
(8)By (7), we have
(9)
Now, we assume that and
, (9) is equivalent to
(10)
Solving in (10),we get that
which implies that
must be the positive root of
Let
where
Since
is a quadratic function in
with
and the coefficient of second order positive, there exists a unique real number
such that
Similarly, the origin is globally asymptotically stable in if
. Otherwise, if
, then
has an equilibrium
with
such that it is globally asymptotically stable in
. Moreover,
is also linearly stable, that is,
has the following from
is stable. The positive components
can be calculated by
where
and
is the unique positive root for
(12)We have the following inequalities
(13)All above computation results will be very useful in the classification for various dynamical behavior. Before finishing this section, we present a result for (3) that is easily obtained by the theory of monotone dynamical systems.
The stability of boundary equilibria
First, in the case that either or
, Theorem 2 tell us that the global behavior for (3) is clear. So it suffices to consider the case both
and
.
From now on, we discuss the stability of the boundary equilibrium .
The Jacobian matrix of (3) at
takes the form
where
is a stable matrix in the above section and
It follows from [27] or Theorem 2.3 in [26] that the stability for the matrices
and
is all the same. By calculation,
(15)From the first equation of
and (6) we get that
(16)and by (7), we have
(17)It deduces from (16) and (17) that
Then, from M-matrix theory [27], it is easy to get that is stable (unstable) if and only if
, that is,
, where
in the above section.
Then we have the results as follows:
Theorem 3.
Let and
.
(I) ,
is stable;
(II) ,
and
,
is stable;
(III) ,
and
,
is unstable.
In a quite similar way, we can discuss the stability for the boundary equilibrium , its stability is completely determined by the determinant of the matrix
The computation shows that
(18)where
is given in (12) and (13).
Observing that and
, we get the following stability results from (18):
Theorem 4.
The stability for
is confirmed by using (18) as follows:
(I) ,
is stable;
(II) ,
and
,
is stable;
(III) ,
and
,
is unstable.
Remark 1.
In Theorem 3 and Theorem 4, we only give the results in this case
. The other cases can be considered analogously by changing the relevant parameters.
Let and
denote the largest real part of its eigenvalues respectively, which is an eigenvalue for
and
respectively by Perron-Frobenius theory [27].
The existence of endemic equilibrium
It follow from Theorem 2 that one of the necessary conditions for existence of positive equilibrium is that and
Now, let we assume is a positive equilibrium for (3), and set
Substituting (22) into (21) yields(23)hence
(24)which implies by
that either
or
In order to study the existence of positive equilibrium, we only need to consider the case (I) and (II). Suppose first the former holds. Without loss of generality, we assume that Let
then
Substituting (25) and (23) into (22), we conclude that such a positive equilibrium must have the form(26)where
Substituting (26) into (21), we obtain the equations for in the form
Then, (27) is reduced to the system(28)
Then, (30) has a unique positive solution if and only if(31)
Moreover, we have the result as follows:
Theorem 5.
If
, and
. System (3) has a unique positive solution if and only if the following conditions is satisfied:
It follows from (4) and Smith [26] that (3) is type-K monotone system, hence tends to an equilibrium as
Then we can give stability conditions for the positive coexistence equilibrium as follows.
Theorem 6.
The positive coexistence equilibrium is stable if
and is unstable if
It remains to consider the case . In this case, it is easy to verify
for
Thus
and
are the same. Let
. Then
. Set
(32)
Then a straight proof by using shows that all points in segment L are nontrivial equilibria for (3).
Theorem 7.
Then nontrivial equilibria set for (3) is L. Moreover, for any
tends to an equilibrium in L as
.
The proof refer to the proof of Theorem 4.2 in [25].
Results
In this article, we have given the stability analysis of the nontrivial boundary equilibria and the positive coexistence equilibrium. Our results can be summarized as the following:
System (3) (and hence (1))has a unique positive coexistence equilibrium if and only if the two nontrivial boundary equilibria have the same stability. (Both are stable or unstable.) The positive coexistence equilibrium is stable if the boundary equilibria are both unstable. In this case the positive coexistence is a globally attractor. The positive coexistence equilibrium is unstable if and only if the boundary equilibria are both stable. The sufficient and necessary conditions for both boundary equilibria to be stable (unstable) and hence for the positive coexistence equilibrium to be unstable (stable) are given by (H1), (H2) in Theorem 5. Furthermore, if there is no coexistence equilibrium, then the locally stable boundary equilibrium, if it exist, is also globally stable.
In the paper [25], we have given the biological meanings for our results. The biological meanings for the results in this paper which can be given in the same way. The interested reader is referred to [25].
Acknowledgments
The authors greatly appreciate an anonymous referee for his valuable comments which improve the paper very much.
Author Contributions
Conceived and designed the experiments: CC JJ. Performed the experiments: CC JJ. Analyzed the data: CC JJ. Contributed reagents/materials/analysis tools: CC JJ. Wrote the paper: CC.
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