1. Introduction
The Allee effect refers to the phenomenon that individuals of many species benefit from the presence of conspecifics, which implies when the per capita growth rate becomes bigger with the increase of the density and gets smaller as long as density passes through a critical value [
1]. When the growth rate decreases but still keeps positive at a low population density, the Allee effect is called weak. It is easy to see that this situation is totally different from the logistic growth. During the last decade this phenomenon attracted a lot of attention with the rise of conservation biology.
The dynamics with the Allee effect, such as the stability of positive equilibrium [
2,
3,
4,
5,
6,
7], the existence of periodic solution [
8,
9,
10,
11,
12], or bifurcation analysis [
13,
14,
15], on nonlinear systems have been extensively investigated. Especially, Boukal and L. Berec [
16] critically reviewed and classified models of single-species population dynamics with the demographic Allee effect. Meanwhile in consideration of the carrying capacity of environment with respect to the prey in the
per capita growth rate of the population, they also proposed the following differential equation with the weak Allee effect
where
denotes the prey population at moment t,
is the
per capita growth rate of the population and
is the carrying capacity of the environment. In biology, the group defense enriches the ability of the prey to defend or escape from the predators, and thus it helps decrease the predation. Tabares et al. [
17] proposed the coupled system of equations
as a model to describe the dynamical behaviour of the predator-prey system incorporating a weak Allee effect in the prey population
and a distributed delay in the predator populations
at time
t. They analyzed the influence of weak Allee effect in the system. Here
and
K are all positive parameters.
is the rate of predation,
is the conversion rate,
is the predator mortality in the absence of prey and
a is the delay parameter.
K denotes the carrying capacity of the environment with respect to the prey. Equation (
2) is regarded as a predator-prey system with group defense [
18]. J. D. Ferreira et al. [
1] proposed the system
which describes the dynamics of interactions between a predator and a prey species with weak Allee effect. They studied the influence of the weak Allee effect with the density function
defined by
and got analogous stability results as (2). For all biological processes in nature, the time delay is an inherent phenomenon. Hence, it is practical to assume that the growth rate of the prey population is determined by its recent history. So taking into account the maturation time of the prey, we define the density function
by
and it makes system (2) the following delayed predator-prey model with weak Allee effect.
In this paper, we study the effect of the delay
on the positive equilibrium of system (4), and investigate the normal form for the Hopf bifurcation. In
Section 2, stability of the equilibrium and existence of Hopf bifurcations are given. In
Section 3, the direction and stability of the bifurcating periodic solutions of the system will be analyzed. Finally, in
Section 4, numerical simulations are performed to illustrate our theoretical results.
2. Stability of the Equilibrium and Existence of Hopf Bifurcations
System (4) can be rewritten as
where
Introducing
,
,
,
and then dropping the bar, system (5) becomes
When
, system (6) is system (3) in [
17]. System (6) has only one positive equilibrium
. In [
17], sufficient conditions, which guarantee the positive equilibrium is locally asymptotically stable for system (6) with
, have been given. We now recall some of their conclusions.
Lemma 1. Equilibrium E of system (6) is locally asymptotically stable provided the following inequalities are satisfied Let
, then Equations (7)–(9) are equivalent to
Denote the three curves on
plane
,
and
by
and
, respectively. For the case when
,
, the three curves divide the first quadrant of the
plane into four regions
(see
Figure 1a). We know from Lemma 1 that the positive equilibrium of system (6) is stable when
.
The characteristic equation of system (6) at E is
i.e.,
Clearly,
is not a root of (13). Let
be a root of (13), then
where
,
,
and
. Separating the real and imaginary parts leads to
It follows directly from (15) that
In fact, (15) can be rewritten as
Now, taking the square of two sides of the equations and adding we obtain (16) directly.
Let
, then (16) can be rewritten as
Since the constant term in (17) is positive,
has at least one positive root if and only if there exists
such that
In fact,
where
and
. When
, it is clearly that
has a positive root
. Through direct calculation, we get
Denote
and
, then curve
divides
into two regions
and
, curve
divides
into two regions
and
(see
Figure 1b). This means that Equation (
17) has two positive roots in region
. Thus, we can draw the following conclusions.
Lemma 2. Assume (10), (11), (18) and (20) hold,
(i) when , has one positive root;
(ii) when , has two positive roots.
A positive root of
corresponds to a pair of purely imaginary roots of Equation (
3). Thus we get the following results.
Lemma 3. Assume (10), (11), (18) and (20) hold.
(i) if has one positive root , then (16) has one pair of purely imaginary roots with double, where ;
(ii) if has two positive roots and , then (16) has two pair of purely imaginary roots and , where and . Furthermore and .
Denote
as a positive root of (16). By (15), we have
Since and , we known .
Since
and
, then
has one positive root and one negative root. Denote the positive root by
. According to the discussions above and Ruan. S et al. [
19], we get the following conclusion.
Lemma 4. (i) If (10), (11), (18) and (20) are not all satisfied, then Equation (13) has at least one root with positive real part for ; (ii) If (10), (11), (20) and hold, then there exists , such that Equation (13) has a pair of purely imaginary roots with double when ; (iii) If (10), (11), (20) and hold, then there exists and such that Equation (13) has a pair of purely imaginary roots when , respectively, whereand Let
be the root of Equation (
13) satisfying
,
, respectively. Substituting
into Equation (
13) and taking derivative with respect to
, we get
where
It follows that
namely,
and
(
), where
. Therefore, the transversality condition holds, and thus Hopf-bifurcation occurs at
. Furthermore, we have the following conclusion.
Lemma 5. Assume (10), (11), (20) and (18) hold.
(i) For , there exists a integer m such that Equation (13) has at least a pair of positive real parts when ; (ii) All roots of Equation (13) have negative real parts when ; Equation (13) has at least a pair of roots with positive real parts when . (iii) when and , all roots of Equation (13) have negative real parts except the purely imaginary roots and . From (24) and (25), it is easy to verify that
Therefore, together with the define of and , means that the lemma is true.
Now we have found the stability switches. In another words, the stability of the equilibrium switching from stability to instability when passes through and back to stability when passes through . When , the equilibrium is instability for ever.
3. Direction and Stability of Hopf Bifurcations
In
Section 2, some conditions which guarantee system (5) and its modification (6) undergo Hopf bifurcation at some critical values of
are obtained. We now apply the normal form and center manifold theory introduced by Hassard et al. [
20] to study the direction of these Hopf bifurcations and stability of the bifurcated periodic solutions.
We rescale the time by
to normalize the delay. Let
and
, then
is the Hopf bifurcation value and system (6) is equivalent to the following functional differential equation in
where
, and
,
are given respectively, by
and
where
,
,
,
,
, and
.
By the Reisz representation theorem, there exist a function
of bounded variation for
such that
In fact, we can choose
where
For
, define
and
Then (28) can be rewritten as
For
, define
and a bilinear inner product
where
. Then
and
are adjoint operators, and
are eigenvalues of
. Of course, they are also eigenvalues of
. Suppose that
is an eigenvector of
corresponding to
, where
. From (31)–(33), we get
Direct computation leads to
It is not difficult to verify that
is an eigenvector of
corresponding to
such that
,
, where
Using the same notations as in literatures [
20,
21,
22,
23], we shall compute the coordinates to describe the center manifold
at
.
Let
be the solution of (28) when
and define
On the center manifold
, we have
where
and
are local coordinates of center manifold
in the direction of the
q and
, respectively. According to the center manifold theory, we have
For solution
in
of Equation (
28),
so we have
Consider the formal Taylor expansion of the
as follows
According to (30), we have
Comparing the coefficients with (43), we have
In order to determine
, we focus on the computation of
and
. By (35), we have
Differentiating (39) with respect to
t, we obtain
Substituting (43) and (46) into (45), and comparing the coefficients, we have
and
According to (33) and (47), for
,
It is clear that the solution of above equation is
For
, we see from (47) that
Substituting (49) into (50), we have
namely,
Thus,
can be determined by the parameters and delay. So the following quantities can be derived:
It is well-known from [
20] that the sign of
decides the direction of Hopf bifurcation, the sign of
determines the stability of bifurcating periodic solutions and
defines the period of bifurcating periodic solutions. Thus we have the following conclusion.
Theorem 1. (1) The Hopf bifurcation is supercritical (subcritical) and the bifurcating periodic solutions exist for if ;
(2) The bifurcating periodic solutions are stable (unstable) if ;
(3) The periodic of the bifurcating periodic solutions increase (decrease) if .
4. Numerical Simulations
In this section, we perform some numerical simulations to verify our theoretical analysis proved in previous sections. To investigate the effect of delay, we focus on the case when . Set , , , , , and . By direct computation, we get and the equilibrium E is .
(i) Take
. Starting from the initial value
, we get
Figure 2. According the previous analysis, the equilibrium should be stable. The simulation results coincide exactly with the our conclusion.
(ii) Take
. Starting from the initial value
, we get
Figure 3. This implies that the Hopf bifurcation associated with the critical value
is supercritical, the bifurcating periodic solution is stable, and the equilibrium E is unstable. All these results are consistent with our analysis.
(iii) Take
. Starting from the initial value
, we get
Figure 4. In this case, the bifurcating periodic solution vanishes, but the equilibrium is still unstable.