1. Introduction
Reaction–diffusion equation models have frequently been used in population dynamics and biological problems. A widely used example is the classical logistic equation,
The spatial domain
is a region in
for
, where
denotes the unit outward normal on the
The variables
t and
x represent time and location separately in the function
of (
1). The function
, under the assumption that
in
, stands for the intrinsic growth rate, and the function
, for all
x in
, stands for the carrying capacity.
Since the carrying capacity of the environment is spatially heterogeneous, i.e.,
K is not a constant function, system (
1) may assume the migration of populations from resource-rich to resource-poor places. To improve on this aspect of diffusion, Korobenko and Braverman [
1] considered the case where an ideal free distribution is a positive state that matches the local per capita growth rate. This led to the following initial boundary value problem:
One can see that the function
is a positive equilibrium solution of system (
2). Moreover, for any
, with
, we have
uniformly present in
. For more details on this, we refer the reader to [
2,
3,
4] and the references therein.
The effect of dispersal on species interaction is a very important topic in spatial ecology. In many cases, the ideal free distribution (IFD), introduced in [
5], is evolutionary stable [
6,
7,
8,
9,
10,
11,
12]. Since most species have a maturation time or a reproduction period, we include a time delay and consider the following model in this present work.
where
represents the time delay, and
with
.
The model above involves heterogeneity in the environment conditions. Hence, the positive equilibrium is spatially non-homogeneous. Moreover, the characteristic equation is no longer an algebraic equation. This makes the study of dynamical behavior near the spatially non-homogeneous positive equilibrium difficult. It is worth noting that after the pioneering work of Busenberg and Huang [
13], many authors investigated the Hopf bifurcation of models with heterogeneity in the environmental conditions [
14,
15,
16,
17,
18,
19,
20,
21,
22,
23,
24,
25,
26].
Dropping the tilde sign, system (
3) becomes
The advantage of this rescale is that the unique positive equilibrium of (
4) becomes equal to 1.
In this paper, we mainly study the influence of the delay
on the stability of the positive equilibrium solution of model (
4). Throughout the paper, we always assume that
and
satisfy the following hypothesis:
Hypothesis 1. 2. Stability and Hopf Bifurcation
Before studying the stability of the delayed nonlinear system (
4), we consider the following eigenvalue problem, which is critical for the determination of the properties of the solutions to the nonlinear Equation (
4):
Due to the Neumann boundary conditions, the principal eigenvalue of (
5) is
. The corresponding eigenfunction (the principal eigenfunction) is strictly positive and unique up to its multiplication by a positive constant. Thus, one may take the principal eigenfunction as
, without loss of generality.
Throughout the paper, we will follow an approach that was recently used in [
27] to study the Hopf bifurcation in a different reaction–advection–diffusion problem with time delay. We let
, and
. Meanwhile,
denotes the complexification of a linear space
,
denotes the domain of a linear operator
L,
is the kernel of
L, and
is the range of
L. Furthermore, we define the inner product of the Hilbert space
as
.
and
denote the Banach space of continuous and differentiable mappings from
into
, respectively.
For convenience of analysis, the spaces
and
are decomposed as
where
The linearization of (
4) at
is given by
We know, from [
28], for example, that the solution semi-group of problem (
6) has an infinitesimal generator
that satisfies
on the domain
where
. From this formulation of (
6), the spectrum of
is
where
Lemma 1. If , then the steady state in (4) is locally asymptotically stable for any . Moreover, for any Proof. When
, computing the spectrum of
leads us to the equation
which in turn leads to the investigation of the following linear eigenvalue problem:
Then,
has the following variational characterization:
This yields . Therefore, the steady state of (3) is locally asymptotically stable when the delay .
We can similarly show for any . □
Next, we prove that the eigenvalues of
will pass through the imaginary axis for some
, which is a necessary condition for the occurrence of the Hopf bifurcation. Indeed,
has a purely imaginary eigenvalue
, for some
, if and only if the equation
is solvable for some values such that
,
,
,
. We are now in place to state the following lemmas.
Lemma 2. If there exist that solve system (9), then is uniformly bounded. Proof. After substituting
into system (
9) and multiplying
, an integration of the result over
leads to
After separating the real and imaginary parts of (
10), we obtain
□
Lemma 3. Let . If and , thenwhere is the second eigenvalue of operator . Proof. Under zero Neumann boundary conditions, it is well-known that the operator
defined on
has a sequence of eigenvalues
, such that
Moreover, the corresponding eigenfunctions
form an orthogonal basis for
. The principal eigenfunction is
In particular, for each
that satisfies
, there is a sequence of real numbers
, such that
. Therefore,
It thus follows from the above equality that
□
Observe that
Hence, if
satisfies system (
9), then we ignore a scalar factor and have
, where
,
and
. By letting
and substituting these into (
9), we obtain
Define from . We confirm in the following lemma that is uniquely solvable when .
Lemma 4. The equationhas a unique solution as . Here, , , and satisfies the following equation Proof. We see from the second equation of (
15) that
(with
) if and only if
. Substituting
into the first equation of (
15), we see that
satisfies
Integrating (
18) over
and then separating the imaginary and real parts, we have
Setting
, we then obtain
□
Next, we prove that there exists , such that has solutions for all .
Lemma 5. There exists and a unique continuously differentiable mapping from to , such that .
Proof. We define
to be the Fréchet derivative of
F with respect to
at the point
, then
We can easily check that T is bijective from to . Thus, by the implicit function theorem, there exists and a continuously differentiable mapping from to , such that .
To prove uniqueness, we appeal to the implicit function theorem, which states that it is sufficient to show that if
then
as
in the norm of
.
We can easily obtain the boundedness of the sequences
,
, and
from the definition, hypothesis, and Lemma 2, respectively. Based on the first equation of (
15) and Lemma 3, we have
The boundedness of
,
, and
implies that there exists a constant
, such that
Therefore, if
is sufficiently small, we have
. Hence,
As a result,
is bounded in
for
. In addition, since the operator
is bounded, we see that
is also bounded in
, which implies that
is pre-compact in
for
. Therefore, there exists a subsequence
, such that
Taking the limit of the equation
as
, we see that
Moreover, since
has a unique solution, we obtain
The proof is completed. □
Remark 1. From Lemma 5, we conclude that for each , the eigenvalue problem (
where , and )
has a solution if and only if Here, .
For convenience, we will always assume in what follows. In fact, considering that further perturbation arguments are used, the interval of might be smaller.
Lemma 6. Assume that . Then, Proof. Since , we have , .
Therefore, . □
Theorem 1. For , there exists a neighborhood of such that has a simple eigenvalue . Moreover, , .
Proof. We know that
, where
. If
, then
Thus, there is a constant
a such that
The first equation of (
22) yields
From (
22) and (
23), we have
By Lemma 6, we obtain that . This implies that is a simple eigenvalue of . By the implicit function theorem, there is a neighborhood of , such that has an eigenvalue for . Moreover, and . □
Now, we study the following transversality condition.
Theorem 2. Assume that . Then, we obtain Proof. Differentiating Equation (
26) with respect to
at
yields
Then, multiplying Equation (
27) by
and integrating it over
, we obtain
This completes the proof. □
On the basis of the lemmas and theorems above, we obtain the following results immediately.
Theorem 3. The infinitesimal generator , , has exactly eigenvalues with positive real parts when , where .
The above results and the theory on partial
functional differential equations, explained in [
28], lead us to the following local Hopf bifurcation theorem.
Theorem 4. For fixed , the positive steady state of (4) is locally asymptotically stable when and is unstable when . Furthermore, system (4) exhibits a Hopf bifurcation at the positive steady state when , . 3. The Direction of the Hopf Bifurcation
The results of
Section 2 demonstrate that periodic solutions bifurcate from the positive steady-state solution
as the delay
passes through the critical value
,
. In this section, with
considered as a bifurcation parameter, we apply the normal form theory and center manifold reduction to analyze the
direction of the Hopf bifurcation that occurs around the positive steady state.
We first transform the steady state
of system (
4) and the critical value
into the origin via the translations
,
and
. Then, dropping the tilde sign, system (
4) becomes
where
Similar to
Section 2, we define that
is the infinitesimal generator of the linearized Equation (
29). Thus, we have
with
where
. Let us also define
Then, (
29) can be rewritten as
It follows from the previous section that has only one pair of simple purely imaginary eigenvalues . The eigenfunction associated with (respectively, ) is (respectively, ) for , where is defined as in Remark 1.
Based on [
27], we introduce the following bilinear form: for
and
,
where
.
The following lemma is concerned with the formal adjoint operator of under the bilinear product above.
Lemma 7. Letfor , where . Then, and satisfy Proof. For
and
, we have
□
Similarly, we see that
has only one pair of simple purely imaginary eigenvalues
. The eigenfunction associated with
(respectively,
) is
(respectively,
) for
where
is defined as in Remark 1. The center subspace of () is
. Furthermore, the basis of the eigenfunction space of the adjoint operator
associated with the eigenvalues
is
. Meanwhile,
is the formal adjoint subspace of
P. As usual,
can be decomposed as follows:
where
One can check that
, where
I is the identity matrix in
. For the bifurcation direction and stability, since the formulas to be built are all only relative to
, we let
in Equation (30) and define
Let
be the center manifold with a range
Q. Then, the flow of (30) on the center manifold can be written as follows:
Since
, we have
where
A straight forward calculation leads to
In order to obtain the stability of bifurcating periodic orbits and the bifurcation directions, we compute the quantities as follows:
and
Motivated by [
28,
29], we have the following results.
Lemma 8. For , Equation (4) creates the Hopf bifurcation at the positive steady state ; moreover, - (i)
determines the direction of the Hopf bifurcation: If (), then the bifurcating periodic solutions exist for (resp. ), and the bifurcation is called forward (resp. backward).
- (ii)
determines the stability of bifurcating periodic solutions: The bifurcating periodic solutions are orbitally asymptotically stable (resp. unstable) if (resp. ).
- (iii)
determines the period of the bifurcating periodic solutions: The period increases (resp. decreases) if (resp. ).
To compute
, we need to figure out
and
. Note that
satisfies
We see that
,
,
, and
satisfy
In what follows, we solve for
E and
F. From (
33) and (
34), together with the definition of
, we see that
E and
F satisfy
Lemma 9. Assume that E and F satisfy (35). Then, , , where and satisfy , , . Moreover, the constant satisfies . Proof. We only give the estimate for
E since the estimate for
F can be obtained similarly. Substituting
into the first equation of system (
35), we get
Multiplying both sides of the Equation (
36) by
and then integrating the result over
,
Multiplying this by
and integrating the result over
, we have
From the lemma above, we then get
Hence, it follows from Equation (
37) that there exist constants
, and
, such that for any
,
Furthermore, from Equations (
38) and (
39), combined with Lemma 3, we obtain that there exist constants
, such that for any
,
Therefore, we have
. This, together with Equation (
37), implies that
□
With arguments similar to those in [
14,
27], one can obtain the following result.
Theorem 5. For , Equation (4) creates the Hopf bifurcation at the positive steady state , near (
)
. Moreover, the direction of the Hopf bifurcation at is forward, and the bifurcating periodic solution from is orbitally asymptotically stable. Proof. Since
,
. Then,
Former conclusions give
and
,
Hence,
We can verify that . This in turn implies that . Using Lemma 8, the proof of this theorem is complete. □