Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

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Commun Nonlinear Sci Numer Simulat

journal homepage: www.elsevier.com/locate/cnsns

Traveling fronts for a delayed reaction–diffusion system with

a quiescent stage
Shi-Liang Wu a,⇑,1, Hai-Qin Zhao b,2
a
Department of Mathematics, Xidian University, Xi’an, Shaanxi 710071, People’s Republic of China
b
Department of Mathematics, Xianyang Normal University, Xianyang, Shaanxi 712000, People’s Republic of China

a r t i c l e i n f o a b s t r a c t

Article history: In this paper, we study the traveling wave fronts of a delayed reaction–diffusion system
Received 21 December 2010 with a quiescent stage for a single species population with two separate mobile and sta-
Received in revised form 8 January 2011 tionary states. By transforming the corresponding wave system into a scalar delayed differ-
Accepted 12 January 2011
ential equation with an integral term, we establish the existence of the minimal wave
Available online 26 January 2011
speed cmin, and the asymptotic behavior, monotonicity and uniqueness (up to a translation)
of the traveling wave fronts. In particular, the effects of the delay and transfer rates on the
Keywords:
minimal wave speed are studied.
Reaction–diffusion system
Traveling wave front
 2011 Elsevier B.V. All rights reserved.
Delay
Quiescent stage

1. Introduction

In recent years, some theory of systems with quiescence has emerged, see e.g., Hadeler et al. [6]. A dynamical system with
known qualitative behavior is interpreted as a system for the evolution of particles, and then diffusively coupled to a
quiescent phase or the zero vector field. The general question is how the dynamics is changed by the introduction of a
quiescent phase. In particular, the classical reaction diffusion equation has been supplied with a quiescent phase, e.g.,
Hadeler and Lewis [5] presented and discussed the following reaction–diffusion system with a quiescent stage

@ t uðx; tÞ ¼ Duxx ðx; tÞ þ f ðuðx; tÞÞ  c1 uðx; tÞ þ c2 v ðx; tÞ;
ð1:1Þ
@ t v ðx; tÞ ¼ c1 uðx; tÞ  c2 v ðx; tÞ;

where u(x, t) and v(x, t) are the densities of mobile and stationary subpopulations at a point x and time t, respectively, D > 0 is
the diffusion coefficient of the mobile subpopulation, c1 > 0 and c2 > 0 are the transfer ratesbetween two states, f(u(x, t)) is the
reproduction function. The system (1.1) could represent a model for a population where individuals migrate and reproduce
and are subject to randomly occurring inactive phases [5].
On the other hand, it is well known that time delays should be taken into consideration in biological models, especially in
those population models, and much has been done in this aspect. Model (1.1) includes a basic assumption: the growth rate of
the population is considered to act instantaneously whereas there may be a time delay to take account of the time of to reach
maturation periods, the hatching period and so on. Thus, as mentioned in [6], it may be worthwhile to study the following
delayed reaction–diffusion system

⇑ Corresponding author.
E-mail address: tunshl03@lzu.cn (S.-L. Wu).
1
Supported by the Fundamental Research Funds for the Central Universities (JY10000970005) and the NSF of China (11026127).
2
Supported by the Talent Project of Xianyang Normal University (06XSYK247).

1007-5704/$ - see front matter  2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.cnsns.2011.01.012
 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621 3611


@ t uðx; tÞ ¼ Duxx ðx; tÞ þ f ðuðx; tÞ; uðx; t  sÞÞ  c1 uðx; tÞ þ c2 v ðx; tÞ;
ð1:2Þ
@ t v ðx; tÞ ¼ c1 uðx; tÞ  c2 v ðx; tÞ;
where the reproduction function is f(u(x, t), u(x, t  s)), s P 0 is a constant, which could be called the natural quiescent exten-
sion of the scalar delayed reaction–diffusion equation
@ t uðx; tÞ ¼ Duxx ðx; tÞ þ f ðuðx; tÞ; uðx; t  sÞÞ:
A special case of system (1.2) is the following delayed diffusive Nicholson’s blowflies equation with a quiescent stage [6]

@ t uðx; tÞ ¼ Duxx ðx; tÞ  dðuðx; tÞÞ þ bðuðx; t  sÞÞel0 s  c1 uðx; tÞ þ c2 v ðx; tÞ;
ð1:3Þ
@ t v ðx; tÞ ¼ c1 uðx; tÞ  c2 v ðx; tÞ;
where u(x, t) and v(x, t) are the densities of mobile and stationary subpopulations of the mature blowflies at a point x and
time t, respectively, l0 is the death rate of the juvenile, the delay s P 0 is the duration of the juvenile state. We refer to
[6] for more details on the model.
Traveling wave solutions are an important class of solutions for a large class of nonlinear reaction–diffusion equations
(systems) modeling a variety of physical and biological phenomena. In the past decades, the study on traveling wave solu-
tions has been extensive and intensive, and has led to many interesting and significant results, see, e.g., [4,10,12,14–16,17–
20] and the surveys of Gourley et al. [3,4]. In [18], Zhang and Zhao considered the asymptotic behavior of system (1.1). More
precisely, they established the existence of spreading speed and showed that it coincides with the minimal wave speed for
monotone traveling wave solutions. Zhang and Li [19] further considered the monotonicity and uniqueness of traveling wave
solutions of (1.1). More recently, Zhao and Wu [21] considered the traveling wave solutions of a lattice differential system
which can be regarded as a spatially discrete version of (1.1) on a 2D spatial lattice.
The purpose of this paper is to study the traveling wave solutions of the delayed reaction–diffusion system (1.2) from the
viewpoint of waves of invasion. In particular, we instigate the effects of the delay and transfer rates on the invasion.
Throughout this paper, we make the following assumptions on the function f:

(A1) f ð0; 0Þ ¼ f ðK; KÞ ¼ 0; f 2 C 2 ð½0; K2 ; RÞ; f ðu; uÞ > 0 for u 2 (0, K), and @ 2f(u, v) P 0 for (u, v) 2 [0, K]2, where K > 0 is a
constant;
(A2) @ 1f(0, 0)u + @ 2f(0, 0)v P f(u, v) for (u, v) 2 [0, K]2, and @ 1f(K, K) + @ 2f(K, K) < 0.

We would like to point out that (A1)–(A2) are standard monostable assumptions, see e.g., [16]. From (A1), it is easy to see
+
that (1.2) has two equilibria E :¼ (0,  0) and E :¼ (K,K0), where K0 = c1K/c2. Furthermore, condition (A1) together with (A2)
imply that @ 1 f ð0; 0Þ þ @ 2 f ð0; 0Þ P K2 f K2 ; K2 > 0. We are interested in traveling wave solutions that connect E and E+. A solu-
tion (u(x, t), v(x, t)) of (1.2) is called a traveling wave solution, if there exist a constant c > 0, and a differential function
ðUðÞ; VðÞÞ : R ! ½0; K  ½0; K 0  such that
ðuðx; tÞ; v ðx; tÞÞ ¼ ðUðx þ ctÞ; Vðx þ ctÞÞ:
The parameter c and (U, V) are called the wave speed and the wave profile, respectively. Moreover, we say (U, V) is a traveling
wave front if (U(), V()) is monotone on R.
We now briefly describe the organization and main ideas of this paper as follows. In Section 2, we first transform the cor-
responding wave system of (1.2) into a scalar delayed differential equation with an integral term. This idea is originated from
Zhao and Wang [20] in studying the traveling waves of a reaction–diffusion system modeling man-environment-man epi-
demics. This property is effectively used to investigate the properties of the traveling wave solutions of (1.2). Then, we estab-
lish the existence of traveling wave fronts for c P cmin :¼ cmin(s, c1, c2) using the classical monotone iteration technique
coupled with the (weak) upper and (weak) lower solutions. Finally, we consider the effects of the delay and transfer rates
c1, c2 on cmin.
In Section 3, we establish the exact asymptotic behavior of the wave profiles as n ? ±1 using Ikehara’s theorem. This
method was developed by Carr and Chmaj [1] for a nonlocal monostable equation, see also [16,19]. As a by-product, the
non-existence of traveling wave solutions for c 2 (0, cmin) is obtained. With these information on the asymptotic behaviors
of traveling wave solutions, we then prove the monotonicity and uniqueness of traveling wave solutions in Section 4 using
a sliding method which was developed by Chen and Guo [2] for a lattice differential equation, see also [19,21]. The paper
ends with a discussion in Section 5.

2. Properties of the minimal wave speed

2.1. Reduction to a scalar equation

Substituting (u(x, t), v(x, t)) = (U(n), V(n)), n = x + ct, into (1.2), we obtain the corresponding wave system
(
cU 0 ðnÞ ¼ DU 00 ðnÞ þ f ðUðnÞ; Uðn  csÞÞ  c1 UðnÞ þ c2 VðnÞ;
ð2:1Þ
cV 0 ðnÞ ¼ c1 UðnÞ  c2 VðnÞ;
3612 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

with the asymptotic boundary condition

ðUð1Þ; Vð1ÞÞ ¼ ð0; 0Þ and ðUðþ1Þ; Vðþ1ÞÞ ¼ ðK; K 0 Þ: ð2:2Þ
We first transform the wave system of (1.2), i.e., (2.1) and (2.2), to a delayed differential equation with an integral term.
From the second equation of (2.1) and V(1) = 0, we have
Z n
c1 c2
VðnÞ ¼ e c ðnsÞ UðsÞds: ð2:3Þ
c 1

Substituting (2.1) into the first equation of (2.1), we get

Z n
c1 c2 c2
cU 0 ðnÞ ¼ DU 00 ðnÞ þ f ðUðnÞ; Uðn  csÞÞ  c1 UðnÞ þ e c ðnsÞ UðsÞds: ð2:4Þ
c 1

Then, the following result holds.

Lemma 2.1

(i) If (U(n), V(n)) is a non-decreasing traveling wave solution of (1.2) , then U(n) is a non-decreasing solution of (2.4);
(ii) If U(n) is a non-decreasing solution of (2.4) with 0 6 UðnÞ 6 K; n 2 R,
Uð1Þ ¼ 0 and Uðþ1Þ ¼ K; ð2:5Þ
and V(n) is defined by (2.3), then (U(n), V(n)) is a non-decreasing traveling wave solution of (1.2).

Consequently, to consider the traveling wave solutions of (1.2), it suffices to study the solutions of problem (2.4) subject
to the boundary condition (2.5).

2.2. Monotone iteration technique

Motivated by the monotone iteration technique in [10,14,17,20], in this subsection, we further reduce the existence of
traveling wave solutions of (1.2) to the construction of an ordered pair of (weak) lower and (upper) solutions of (2.4).
Let C ½0;K ðR; RÞ ¼ f/ 2 CðR; RÞ : 0 6 /ðsÞ 6 K; s 2 Rg: Define the operator H on C ½0;K ðR; RÞ by
Z n
c1 c2 c2
Hð/ÞðnÞ ¼ f ð/ðnÞ; /ðn  csÞÞ þ L1 /ðnÞ þ e c ðnsÞ /ðsÞds; ð2:6Þ
c 1

where L1 ¼ maxðu;v Þ2½0;K2 fj@ 1 f ðu; v Þjg. With the notation, (2.4) is equivalent to the following ordinary differential equation
DU 00 ðnÞ  cU 0 ðnÞ  ðc1 þ L1 ÞUðnÞ þ HðUÞðnÞ ¼ 0:
Moreover, one can easily verify that H possesses the following properties.

Lemma 2.2. Assume that (A1) holds. Then

(i) H(/1)(n) 6 H(/2)(n), for all /1 ; /2 2 C ½0;K ðR; RÞ with /1 6 /2;

(ii) H(/)(n) is non-decreasing in n 2 R, if / 2 C ½0;K ðR; RÞ is non-decreasing in n 2 R.

Define F : C ½0;K ðR; RÞ ! CðR; RÞ by

Z n Z þ1 
1
Fð/ÞðnÞ ¼ ek1 ðnsÞ Hð/ÞðsÞds þ ek2 ðnsÞ Hð/ÞðsÞds ; ð2:7Þ
Dðk2  k1 Þ 1 n

where
pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
c c2 þ 4ðc1 þ L1 Þ c þ c2 þ 4ðc1 þ L1 Þ
k1 ¼ and k2 ¼ :
2D 2D
Clearly, F is well defined on C ½0;K ðR; RÞ, and for any / 2 C ½0;K ðR; RÞ; Fð/Þ satisfies
DðFð/ÞÞ00 ðnÞ  cðFð/ÞÞ0 ðnÞ  ðc1 þ L1 ÞFð/ÞðnÞ þ Hð/ÞðnÞ ¼ 0: ð2:8Þ
Thus, a fixed point of F is a solution of (2.4), which is a traveling wave solution of (1.2).
The following result follows from Lemma 2.2, see e.g., Wu and Zou [17] and Huang and Zou [7].

Lemma 2.3. Assume that (A1) holds. Then

(i) F(/1)(n) 6 F(/2)(n), for all /1 ; /2 2 C ½0;K ðR; RÞ with /1 6 /2;

(ii) F(/)(n) is non-decreasing in n 2 R, if / 2 C ½0;K ðR; RÞ is non-decreasing in n 2 R.
 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621 3613

In order to use the method of upper and lower solutions, we introduce the following definition.

 and / are called an upper and a lower solution of (2.4), respectively, if

Definition 2.4. A pair of continuous functions /
 / 2 C 2 ðR; RÞ, and /;
/;  / satisfy
Z n
 0 ðnÞ P D/
 00 ðnÞ þ f ð/ðnÞ;
 
  csÞÞ  c /ðnÞ cc c2

c/ /ðn 1 þ 1 2 e c ðnsÞ /ðsÞds; ð2:9Þ
c 1

and
Z n
c1 c2 c2
c/0 ðnÞ 6 D/00 ðnÞ þ f ð/ðnÞ; /ðn  csÞÞ  c1 /ðnÞ þ e c ðnsÞ /ðsÞds: ð2:10Þ
c 1


Proposition 2.5. Assume that (A1) holds, and that (2.4) admits an upper solution /ðnÞ and a lower solution /(n) satisfying

 6 K; /ðnÞ
(H) 0 6 /ðÞ 6 /ðÞ  
is non-decreasing on R; /ð1Þ 
¼ 0; /ðþ1Þ ¼ K, and / X 0.Then (1.2) has a non-decreasing trav-
eling wave solution (U(n), V(n)).

Proof. By using the classical monotone iteration technique coupled with the upper and lower solutions, the assertion can be
verified as in the proofs of Zhao and Wang [20, Theorem2.1] and Wu and Zou [17, Theorem 3.6]. We only sketch the outline.
Consider the following iteration scheme:
(

/0 ðnÞ ¼ /ðnÞ;
/nþ1 ðnÞ ¼ Fð/n ÞðnÞ; n ¼ 1; 2; . . . :

Using the properties of F and the definitions of the upper and lower solutions, one can show that /n(n) is non-decreasing on R

and /ðnÞ 6 /nþ1 ðnÞ 6 /n ðnÞ 6 /ðnÞ; 
n 2 R. Thus, U(n) = limn?1/n(n) exists and satisfies /ðnÞ 6 UðnÞ 6 /ðnÞ; n 2 R. Moreover, it
is easy to see that U(1) = 0 and U(+1) = K. Define
Z n
c1 c2
VðnÞ ¼ e c ðnsÞ UðsÞds;
c 1

it follows that (U(n), V(n)) is a monotone traveling wave solution of (1.2). This completes the proof. h

The upper and lower solutions in Proposition 2.5 are difficult to construct, since they are required to be twice continu-
ously differentiable on R. In order to overcome the difficulty, similar to Ma [10] and Li et al. [9], we introduce the following
weak upper solution and weak lower solution of (2.4), which is easy to construct in practice.

Definition 2.6. A continuous functions /  is called a weak upper solution of (2.4), if there exist Ti, i = 1,. . .,m such that /
 are
 satisfies
twice continuously differentiable in R n fT i : i ¼ 1; . . . ; mg and /
Z n
 0 ðnÞ P D/
 00 ðnÞ þ f ð/ðnÞ;
 
  csÞÞ  c /ðnÞ cc c2

c/ /ðn 1 þ 1 2 e c ðnsÞ /ðsÞds; ð2:11Þ
c 1

for n 2 R n fT i : i ¼ 1; . . . ; mg. A weak lower solution of (2.4) can be defined similarly.

 / 2 C ½0;K ðR; R2 Þ are a weak upper solution and a weak lower solution of (2.4) , respec-
Lemma 2.7. Assume that (A1) holds. If /;
tively, satisfying (H) and
 0 ðnþÞ 6 /
/  0 ðnÞ; /0 ðnþÞ P /0 ðnÞ; ð2:12Þ

then, Fð/Þð1Þ 
¼ 0; Fð/Þðþ1Þ ¼ K,

/ðnÞ 6 Fð/ÞðnÞ 6 Fð/ÞðnÞ 
6 /ðnÞ; n 2 R; ð2:13Þ

and q  q :¼ Fð/Þ 2 C ½0;K ðR; RÞ are an upper solution and a lower solution of (2.4), respectively.
 :¼ Fð/Þ;

Proof. The proof is similar to those of [9, Lemma 3.9] and [10, Lemmas 2.5-2.6]. We omit it here. h

The following result follows from Lemma 2.7 and Proposition 2.5.

Proposition 2.8. Assume that (A1) holds, and that (2.4) has a weak upper solution / and a weak lower solution / satisfying (H)
and (2.12) , then (1.2) has a non-decreasing traveling wave solution (U(n), V(n)).
3614 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

2.3. Existence of a minimal wave speed

For c P 0 and k 2 C with k–  cc2 , we define two functions:

c1 c2
D1 ðc; kÞ ¼ ck  Dk2  @ 1 f ð0; 0Þ  @ 2 f ð0; 0Þekcs þ c1  ;
ck þ c2
c1 c2
D2 ðc; kÞ ¼ ck  Dk2  @ 1 f ðK; KÞ  @ 2 f ðK; KÞekcs þ c1  :
ck þ c2
It is easy to see that the following result holds.

Lemma 2.9. Assume that @ 1f(0, 0) + @ 2f(0, 0) > 0, o1f(K, K) + @ 2f(K, K) < 0, and c1 P 0, c2 > 0. Then,

(i) there exists a positive number cmin :¼ cmin(s, c1, c2) such that
(a) if 0 6 c < cmin and k P 0, then D1(c, k) < 0;
(b) if c P cmin, then the equation D1(c, k) = 0 has two positive real roots k1(c) and k2(c) with k1(c) 6 k2(c);
(c) if c = cmin, then k1(cmin) = k2(cmin) :¼ k⁄, and if c > cmin, then k1 ðcÞ < k < k2 ðcÞ; k01 ðcÞ < 0; k02 ðcÞ > 0, and
D1 ðc; Þ < 0 in R n ðk1 ðcÞ; k2 ðcÞÞ; D1 ðc; Þ > 0 in ðk1 ðcÞ; k2 ðcÞÞ:
(ii) the equation D2(c, k) = 0 has two real roots k3(c) < 0 and k4(c) > 0.

Lemma
 2.10. Assume that (A1) and (A2) hold. Let c > cmin and k1(c), k2(c) be defined as in Lemma 2.9. Then for every
b 2 1; minf2; kk21 ðcÞ
ðcÞ
g , there exists Q(c, b) P 1, such that for any q P Q(c, b), the function /± defined by


/þ ðnÞ ¼ min K; ek1 ðcÞn þ qebk1 ðcÞn ; n 2 R;
and


/ ðnÞ ¼ max 0; ek1 ðcÞn  qebk1 ðcÞn ; n 2 R;
are a weak upper solution and a weak lower solution of (2.4), respectively.

Proof. By virtue of f 2 C 2 ð½0; K2 ; RÞ and @ 1f(0, 0)u + @ 2f(0, 0)v P f(u, v) for (u, v) 2 [0, K]2, one can easily verify that /+(n) and
/(n) are a weak upper solution and a weak lower solution of (2.4), respectively. We omit the details. h

Proposition 2.11. Assume that (A1) and (A2) hold. Then there exists a number cmin > 0 ( given by Lemma 2.9 ) such that the fol-
lowing results hold:

(i) for every c P cmin, (1.2) has a traveling wave front (U(n), V(n)) with speed c.
(ii) for 0 < c < cmin, (1.2) has no traveling wave solutions.

Proof

(i) . For c > cmin, the existence of traveling wave fronts follows from Proposition 2.8 and Lemma 2.10. For c = cmin, the exis-
tence of traveling wave fronts can be proved by a limiting argument similar to that of [20, Theorem3.1]. We omit the
details.
(ii) The non-existence of traveling wave solutions will be proved in Section 3. This completes the proof. h

2.4. The effects of delay and transfer rates on cmin

In this subsection, we shall consider the effects of the delay s and transfer rates c1, c2 on the minimal wave speed
cmin(s, c1, c2).
From Lemma 2.9, we know that

@ 
D1 ðcmin ; k Þ ¼ 0 and D1 ðc ; kÞ ¼ 0; ð2:14Þ
@k
min 
k¼k

Fixed c1, c2, it then follows from (2.14) that cmin = cmin(s) and k⁄ = k⁄(s) are differentiable functions with respect to s, and

 
dcmin @ D1 @ D1  ck@ 2 f ð0; 0Þekcs 

¼ ¼ 6 0;
ds @s @c ðc min ;k Þ
k þ ks@ 2 f ð0; 0Þekc s c1 c2 k 
þ ðckþc Þ2 
2 ðcmin ;k Þ
 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621 3615

which implies that

(i) if @ 2f(0, 0) > 0, then the delay will slow the minimal wave speed;
(ii) if @ 2f(0, 0) = 0, then the delay is independent of the minimal wave speed. It should be mentioned that some similar
results have been obtained in Wang et al. [16] for a spatial continuous delayed nonlocal reaction–diffusion equation.

Next, we consider the effect of c1, c2 on cmin. Similarly, we have


 ck 
dcmin @ D1 @ D1  ckþc2 

¼  ¼ c1 c2 k  < 0;
dc1 @ c1 @c ðcmin ;k Þ k þ ks@ 2 f ð0; 0Þekcs þ 
ðckþc Þ2
2 ðcmin ;k Þ

and 
 cc1 k 
dcmin @ D1 @ D1  ðckþc2 Þ2 

¼ ¼ > 0:
dc2 @ c2 @c ðcmin ;k Þ k þ ks@ 2 f ð0; 0Þekcs þ c1 c2 k 2 
ðckþc Þ 2 ðcmin ;k Þ

Thus, cmin is a decreasing function of c1 and an increasing function of c2. This finding agrees with physical intuition: increas-
ing the sojourn time in the quiescent phase decreases the propagation speed.
Let us now consider an important cases, namely c1 = 0. In this case, there are no individuals switch from mobile state to
stationary state. It is easy to see that cmin equals the minimal wave speed c1 of the Fisher-KPP equation without a quiescent
stage
@ t uðx; tÞ ¼ Duxx ðx; tÞ þ f ðuðx; tÞ; uðx; t  sÞÞ; ð2:15Þ
that is, c1 satisfies

@ 
D0 ðc1 ; k1 Þ ¼ 0 and D0 ðc1 ; kÞ ¼ 0;
@k k¼k1

where
D0 ðc; kÞ ¼ ck  Dk2  @ 1 f ð0; 0Þ  @ 2 f ð0; 0Þekcs :
This model (2.15) assumes that individuals are always mobile.
Moreover, one can see that.

(i) cmin(s, c1, c2) > 0 for any s P 0 and c1, c2 > 0, which implies that the invasion can occur even when the transfer rates are
infinitesimally small;
(ii) cmin ðs; c1 ; c2 Þ < c1 ðsÞ for any s P 0 and c1, c2 > 0. Thus, for population switching between mobile and stationary states,
Fisher’s formula of spread rates will overestimate their true value.

3. Asymptotic behavior of traveling waves

To prove the the asymptotic behavior of the wave profiles, the following lemma is important, which can be found in Carr
and Chmaj [1].
R þ1
Lemma 3.1. Let FðKÞ :¼ 0 eKn uðnÞdn and u(n) be a positive decreasing function. If F can be written as F(K) = J(K)
(k+1)
(K + K0) for some constants k > 1, K0 > 0, and some analytic function J in the strip  K0 6 ReK < 0, then
uðnÞ JðK0 Þ
lim ¼ :
n!þ1 nk eK0 n CðK0 þ 1Þ

Lemma 3.2. Let (U(n), V(n)) be a traveling wave solution of (1.2). Then

(i) U(n) 2 (0, K) and VðnÞ 2 ð0; K 0 Þ; n 2 R;

(ii) limn?1U0 (n) = limn? 1V0 (n) = 0.

Proof

(i) By virtue of 0 6 UðnÞ 6 K; n 2 R and U(n) = F(U)(n), i.e.,

Z n Z þ1 
1
UðnÞ ¼ ek1 ðnsÞ HðUÞðsÞds þ ek2 ðnsÞ HðUÞðsÞds ; ð3:1Þ
Dðk2  k1 Þ 1 n
3616 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

it is easy to show that UðnÞ 2 ð0; KÞ; n 2 R. It then follows from

Z n
c1 c2
VðnÞ ¼ e c ðnsÞ UðsÞds
c 1

that VðnÞ 2 ð0; K 0 Þ; n 2 R.

(ii) Since limn?1U(n) = limn?1V(n) = 0,
c1 c2
lim V 0 ðnÞ ¼ lim UðnÞ  lim VðnÞ ¼ 0:
n!1 c n!1 c n!1

From (3.1), we have

Z n Z þ1 
1
U 0 ðnÞ ¼ k1 ek1 ðnsÞ HðUÞðsÞds þ k2 ek2 ðnsÞ HðUÞðsÞds :
Dðk2  k1 Þ 1 n

Then, using the L. Hospital’s rule, it is easy to verify that limn?1U0 (n) = 0. This completes the proof. h

Proposition 3.3 (Asymptotic behavior). Assume that (A1)–(A2) hold. Let (U(n), V(n)) be a traveling wave solution of (1.2) with
speed c P cmin. Then the following hold:

(i) for c > cmin,

lim UðnÞek1 ðcÞn ¼ a0 ðcÞ; lim U 0 ðnÞek1 ðcÞn ¼ a0 ðcÞk1 ðcÞ; ð3:2Þ
n!1 n!1

lim VðnÞek1 ðcÞn ¼ b0 ðcÞa0 ðcÞ; lim V 0 ðnÞek1 ðcÞn ¼ b0 ðcÞa0 ðcÞk1 ðcÞ; ð3:3Þ
n!1 n!1

and for c = cmin,

lim UðnÞn1 ek1 ðcÞn ¼ a0 ðcÞ; lim U 0 ðnÞn1 ek1 ðcÞn ¼ a0 ðcÞk1 ðcÞ; ð3:4Þ
n!1 n!1
0
lim VðnÞn1 ek1 ðcÞn ¼ b0 ðcÞa0 ðcÞ; lim V ðnÞn1 ek1 ðcÞn ¼ b0 ðcÞa0 ðcÞk1 ðcÞ; ð3:5Þ
n!1 n!1

(ii) for c P cmin,

lim ðK  UðnÞÞek3 ðcÞn ¼ a1 ðcÞ; lim U 0 ðnÞek3 ðcÞn ¼ a1 ðcÞk3 ðcÞ; ð3:6Þ
n!þ1 n!þ1

c1 a1 ðcÞ c1 a1 ðcÞk3 ðcÞ

lim ðK 0  VðnÞÞek3 ðcÞn ¼ ; lim V 0 ðnÞek3 ðcÞn ¼  ; ð3:7Þ
n!þ1 ck3 ðcÞ þ c2 n!þ1 ck3 ðcÞ þ c2
c1
where a0(c), a1(c) are constants, b0 ðcÞ ¼ ck1 ðcÞþc ; k1 ðcÞ and k3(c) are defined as in Lemma 2.9.
2

Proof

(i) Note that V() satisfies

Z n
c1 c2
VðnÞ ¼ e c ðnsÞ UðsÞds:
c 1

To complete the proof of the conclusion (i), it suffices to show that (3.2) and (3.4) hold. We divide the proof into three
steps.
Step 1. We show that U(n) is integrable on (1, n0 ] for some n0 2 R. Recall that U satisfies the Eq. (2.4). Denote
q1 = @ 1f(0, 0) + @ 2f(0, 0) and q2 = @ 2f(0, 0)  @ 1f(0, 0). Note that q1 > 0 and U(1) = 0. There exists n0 < 0 such that
for n 6 n0 ,
q1 q1
L½U 2 ðnÞ þ 2UðnÞUðn  csÞ þ U 2 ðn  csÞ 6 UðnÞ þ Uðn  csÞ:
4 4
Thus, by Taylor’s expansion, for n 6 n0 ,

f ðUðnÞ; Uðn  csÞÞ P @ 1 f ð0; 0ÞUðnÞ þ @ 2 f ð0; 0ÞUðn  csÞ  L½U 2 ðnÞ þ 2UðnÞUðn  csÞ þ U 2 ðn  csÞ
q1 q2
P UðnÞ þ ½Uðn  csÞ  UðnÞ;
4 2
Take
Z n Z 0
e c c2 c2 c2
UðnÞ :¼ 2 e c ðnsÞ UðsÞds ¼ e c s Uðs þ nÞds:
c 1 c 1
 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621 3617

It then follows from (2.4) that for n 6 n0 ,

q1 q2 e
cU 0 ðnÞ P DU 00 ðnÞ þ UðnÞ þ ½Uðn  csÞ  UðnÞ þ c1 ½ UðnÞ  UðnÞ: ð3:8Þ
4 2
Integrating (3.8) over [y, n], y < n 6 n0 , we obtain
Z n Z n Z n
q1 q2 e
c½UðnÞ  UðyÞ P D½U 0 ðnÞ  U 0 ðyÞ þ UðsÞds þ ½Uðs  csÞ  UðsÞds þ c1 ½ UðsÞ  UðsÞds: ð3:9Þ
4 y 2 y y

Note that U(1) = 0, U0 (1) = 0,

Z n Z 0 Z 1 Z 0 Z 1
e c2 c2 c2 c2
lim ½ UðsÞ  UðsÞds ¼ lim re c r ½Uðn þ hrÞ  Uðy þ hrÞdhdr ¼ re c r Uðn þ hrÞdhdr; ð3:10Þ
y!1 y c y!1 1 0 c 1 0

and
Z n Z 1
lim ½Uðs  csÞ  UðsÞds ¼ cs Uðn  hcsÞdh: ð3:11Þ
y!1 y 0

Then, letting y ? 1 in (3.9), we obtain

Z n Z 1 Z 0 Z 1
q1 csq2 c1 c2 c2
cUðnÞ P DU 0 ðnÞ þ UðsÞds  Uðn  hcsÞdh þ re c r Uðn þ hrÞdhdr; ð3:12Þ
4 1 2 0 c 1 0

e
which implies that U(n) is integrable on (1, n0 ]. It then follows from (3.8) that UðnÞ is integrable on (1, n0 ].
Rn
Step 2. We show that U(n) = O(ecn) as n ? 1 for some c > 0. Define a function WðnÞ ¼ 1 UðsÞds; n 6 n0 , which is increas-
ing and satisfies W(1) = 0. We claim that W(n) = O(ecn) as n ? 1 for some c > 0.
Integrating (3.8) form 1 to n, n 6 n0 , we obtain
Z n 
q1 q2 e
cUðnÞ P DU 0 ðnÞ þ WðnÞ þ ½Wðn  csÞ  WðnÞ þ c1 UðsÞds  WðnÞ :
4 2 1
Rn R1
Note that y
½Wðs  csÞ  WðsÞds ! cs 0
Wðn  hcsÞdh as y ! 1; and
Z n Z n Z 0 Z 0
e c c2 c2 c2
UðsÞds ¼ 2 lim e c r Uðr þ sÞdrds ¼ e c r Wðn þ rÞdr;
1 c y!1 y 1 c 1

and
Z n Z 0 Z 0 Z 1
c2 c2
lim e c r ½Wðs þ rÞ  WðsÞdrds ¼ re c r Wðn þ hrÞdhdr;
y!1 y 1 1 0

we obtain
Z 1 Z n Z 0 Z 1
csjq2 j q1 c1 c2 c2
cWðnÞ þ Wðn  hcsÞdh P DUðnÞ þ WðsÞds þ re c r Wðn þ hrÞdhdr: ð3:13Þ
2 0 4 1 c 1 0

Since W is increasing, we have

 Z  Z Z
c c þ1 cc2 r csjq2 j q n q 0
cþ 1 2 re dr þ WðnÞ P 1 WðsÞds ¼ 1 Wðn þ sÞds:
c 0 2 4 1 4 1
Hence, for any l > 0 and n 6 n0 ,
  Z
c sjq2 j q 0 q
c 1þ 1 þ WðnÞ P 1 Wðn þ sÞds P 1 lWðn  lÞ:
c2 2 4 l 4
Choose l0 > 0 sufficiently large such that h0 :¼ c(4c1 + 4c2 + 2sjq2jc2)/(q1 l0c2) 2 (0, 1). Thus, W(n  l0) 6 h0W(n), "n 6 n0 . De-
f ðnÞ ¼ WðnÞecn , where c ¼ 1 ln 1 . Then for any n 6 n0 ; W
fine W f ðn  l0 Þ 6 W
f ðnÞ: Hence, 0 6 W f ðnÞ 6 K 0 :¼ maxf W
f ðsÞjs 2
l0 h0
½n0  l0 ; n0 g, for n 6 n0 , which implies that W(n) = O(ecn), as n ? 1. From (3.13), we have U(n) = O(ecn), as n ? 1.
Step 3. For 0Z<þ1 Rek < c, we define a two-sided Laplace transform of U by
LðkÞ ¼ UðnÞekn dn:
1

Rewrite (2.4) as
Z n
c1 c2 c2
cU 0 ðnÞ  DU 00 ðnÞ  @ 1 f ð0; 0ÞUðnÞ  @ 2 f ð0; 0ÞUðn  csÞ þ c1 UðnÞ  e c ðnsÞ UðsÞds
c 1

¼ f ðUðnÞ; Uðn  csÞÞ  @ 1 f ð0; 0ÞUðnÞ  @ 2 f ð0; 0ÞUðn  csÞ: ð3:14Þ

3618 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

Multiplying the two sides of (3.14) by ekn and integrating along n on R, we get
Z þ1
D1 ðc; kÞLðkÞ ¼ ekn ½f ðUðnÞ; Uðn  csÞÞ  @ 1 f ð0; 0ÞUðnÞ  @ 2 f ð0; 0ÞUðn  csÞdn: ð3:15Þ
1

Let u(n) = U(n) and K = k in (3.15), then u(+1) = 0 and

Z þ1
D1 ðc; KÞL1 ðKÞ ¼ eKn ½f ðuðnÞ; uðn þ csÞÞ  @ 1 f ð0; 0ÞuðnÞ  @ 2 f ð0; 0Þuðn þ csÞdn; ð3:16Þ
1
R þ1
where L1 ðKÞ ¼ 1 uðnÞeKn dn:
By virtue of Lemma 3.1, the remainder of the proof is similar to that of [16, Theorem 4.8], and is omitted here.
(ii) Let Z(n) = K  U(n) and F(u,v) = f(K  u,K  v), then Z(+1) = 0,0 < Z() < K,

@ 1 Fð0; 0Þ þ @ 2 Fð0; 0Þ ¼ @ 1 f ðK; KÞ  @ 2 f ðK; KÞ > 0;

and Z(n) satisfies
Z n
c1 c2 c2
cZ 0 ðnÞ ¼ DZ 00 ðnÞ  FðZðnÞ; Zðn  csÞÞ  c1 ZðnÞ þ e c ðnsÞ ZðsÞds:
c 1

Note that for c P cmin and k3(c) is a single real root of the equation D2(c, k) = 0. Using the above argument, the conclusion can
be shown by a little modification. This completes the proof. h

Corollary 3.4. Let the assumptions of Proposition 3.3 be satisfied. Then, for all c P cmin,
U 0 ðnÞ V 0 ðnÞ U 0 ðnÞ V 0 ðnÞ
lim ¼ lim ¼ k1 ðcÞ and lim ¼ lim ¼ k3 ðcÞ:
n!1 UðnÞ n!1 VðnÞ n!þ1 UðnÞ  K n!þ1 VðnÞ  K 0

Proof of Proposition 2.11 (ii). We now show that for 0 < c < cmin, (1.2) has no traveling wave solution. Suppose for the con-
trary that (1.2) has a traveling wave solution. Then (2.4) has a solution U(n) satisfies U(1) = 0 and U(+1) = K. Form Lemma
2.9, D1(c, K) has no real zeroes, and hence it follows from (3.16) that L1 ðKÞ is defined for K with ReK < 0. Moreover,
Z þ1
eKn ½D1 ðc; KÞuðnÞ  f ðuðnÞ; uðn þ csÞÞ þ @ 1 f ð0; 0ÞuðnÞ þ @ 2 f ð0; 0Þuðn þ csÞdn ¼ 0:
1

Since limK?1 D1(c, K) = 1, we obtain a contradiction, and the assertion of the conclusion follows.

4. Monotonicity and uniqueness of traveling waves

We have the following results on the monotonicity and uniqueness of traveling wave solutions of (1.2).

Proposition 4.1 (Monotonicity). Let (U(n), V(n)) be a traveling wave solution of (1.2) with speed c P cmin. Then U0 (n) > 0 and
V 0 ðnÞ > 0; n 2 R.

Proposition 4.2 (Uniqueness). Let (U1(n), V1(n)) and (U2(n), V2(n)) be two traveling wave solutions of (1.2) with speed c P cmin.
Then there exists n0 2 R such that
ðU 1 ð þ n0 Þ; V 1 ð þ n0 ÞÞ  ðU 2 ðÞ; V 2 ðÞÞ:
To prove the two Propositions, we need a few lemmas.

Lemma 4.3. Let (U(n), V(n)) be a traveling wave solution of (1.2). Then the following statements hold:

(i) If U 0 ðnÞ P 0; n 2 R, then U 0 ðnÞ > 0; n 2 R;

(ii) If U 0 ðnÞ > 0; n 2 R, then V 0 ðnÞ > 0; n 2 R.

Proof. Let W(n) = U0 (n), then

cW 0 ðnÞ ¼ DW 00 ðnÞ þ @ 1 f ðUðnÞ; Uðn  csÞÞWðnÞ þ @ 2 f ðUðnÞ; Uðn  csÞÞWðn  csÞ  c1 WðnÞ þ c2 V 0 ðnÞ:

Suppose for the contradiction that there exists n1 2 R such that U0 (n1) = 0, i.e., W(n1) = 0. Since W(n) = U0 (n) P 0,W0 (n1) = 0,
W00 (n1) P 0, and

0 P @ 2 f ðUðn1 Þ; Uðn1  csÞÞWðn1  csÞ þ c2 V 0 ðn1 Þ: ð4:1Þ

 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621 3619

Rn c2
Since U0 (n) P 0 and VðnÞ ¼ cc1 1 e c ðnsÞ UðsÞds, one can easily see that V0 (n) P 0. Note that @ 2f(u, v) P 0 for (u, v) 2 [0, K]2, it
follows from (4.1) that 0 P c2V (n1) P 0, and hence V0 (n1) = 0. By direct computation, we have
0

Z n1
c1 c2
0 ¼ V 0 ðn1 Þ ¼ e c ðn1 sÞ ½Uðn1 Þ  UðsÞds P 0;
c 1

so, U(n1) = U(s) for s 6 n1. Let s ? 1, we get U(n1) = 0, which contradicts the fact that U(n) 2 (0, K). This proves the assertion
(i). The assertion (ii) follows from the equality (2.3), i.e.,
Z n
c1 c2
VðnÞ ¼ e c ðnsÞ UðsÞds:
c 1

This completes the proof. h

Lemma 4.4. Let (Ui(n), Vi(n)) be a traveling wave solution of (1.2) , i = 1, 2. If there exists n 2 R such that U1(+n⁄)  U2(), then
V1(+n⁄)  V2().

Proof. The assertion follows from the equality (2.3), and is omitted. h

From Lemmas 4.3 and 4.4, to prove the monotonicity and uniqueness of traveling wave solutions of (1.2), i.e., Propositions
4.1,4.2, it suffices to show that the solutions of (2.4), (2.5) are monotone and unique.
For this, we first establish a strong comparison theorem for the solutions of (2.4), (2.5).

Lemma 4.5. Let Ui(n) be a solution of (2.4), (2.5) with 0 6 U i ðnÞ 6 K; n 2 R; i ¼ 1; 2; and satisfying U1 P U2 on R. Then either
U1  U2 or U1 > U2 on R.

Proof. Suppose that there exists n2 2 R such that U1(n2) = U2(n2). Then,
Z n2 Z þ1
0 ¼ Dðk2  k1 Þ½U 1 ðn2 Þ  U 2 ðn2 Þ ¼ ek1 ðn2 sÞ ½HðUÞðsÞ  HðU 2 ÞðsÞds þ ek2 ðn2 sÞ ½HðUÞðsÞ  HðU 2 ÞðsÞds
1 n2

Since H[U1]() P H[U2](), H[U1](n) = H[U2](n) for n 2 R. Note that @ 2f(u, v) P 0 for (u, v) 2 [0, K]2. Thus, for any n 2 R,
0 ¼ H½U 1 ðnÞ  H½U 2 ðnÞ
Z n
c1 c2 c2
¼ f ðU 1 ðnÞ; U 1 ðn  csÞÞ  f ðU 2 ðnÞ; U 2 ðn  csÞÞ þ L1 ½U 1 ðnÞÞ  U 2 ðnÞ þ e c ðnsÞ ½U 1 ðsÞ  U 2 ðsÞds
c 1
Z n
c1 c2 c2
P e c ðnsÞ ½U 1 ðsÞ  U 2 ðsÞds P 0;
c 1

it follows that U1(n) = U2(n) for any n 2 R, i.e., U1  U2. The proof is complete. h

By virtue of the strong comparison theorem, one can prove the monotonicity and uniqueness of solutions of (2.4), (2.5) by
using the sliding method as in[2]. Hence, we only sketch the outline.
In what follows, we always assume the condition (A1) and (A2) hold.

Lemma 4.6. Let U(n) be a solution of (2.4), (2.5) with 0 6 UðnÞ 6 K; n 2 R. Then U 0 ðnÞ > 0; n 2 R.

Proof. From Corollary 3.4, we know that U(n) is strictly increasing in R n ½M; M for some M  1. By Lemma 4.5 and using
the sliding method similar to that of [2, Lemma 4.3], one can easily show that U 0 ðnÞ > 0; n 2 R. This completes the proof. h

Lemma 4.7. Let U(n) be a solution of (2.4) with 0 6 UðnÞ 6 K; n 2 R. Then there exists q0 = q0(c, f) 2 (0, 1) such that for any
q 2 (0, q0],
ð1 þ qÞf ðUðnÞ; Uðn  csÞÞ  f ðð1 þ qÞUðnÞ; ð1 þ qÞUðn  csÞÞ > 0;
on {njU(n) > K  q0}.
For a given solution U(n) of (2.4), (2.5) with 0 6 UðnÞ 6 K; n 2 R, we define
  
UðnÞ 
j ¼ jðUÞ :¼ sup UðnÞ 6 K  q0 :
U 0 ðnÞ
Clearly, 0 < j < +1, since limn?1U0 (n)/U(n) = k1(c) > 0 and U0 () > 0 on R.
3620 S.-L. Wu, H.-Q. Zhao / Commun Nonlinear Sci Numer Simulat 16 (2011) 3610–3621

The following result follows from Lemmas 4.6 and 4.7.

Lemma 4.8. Let Ui(n) be a solution of (2.4), (2.5) with 0 6 U i ðnÞ 6 K; n 2 R; i ¼ 1; 2. Assume that there exists q 2 (0, q0] such that
(1 + q)U1(  jq) P U2() on R, where j = j(U1). Then U1() P U2() in R.

Lemma 4.9. Let Ui(n) be a solution of (2.4), (2.5) with 0 6 U i ðnÞ 6 K; n 2 R; i ¼ 1; 2. Then there exists n0 2 R such that U1(+n0) 
U2().

Proof. By virtue of Lemmas 4.5 and 4.8 and applying the sliding method, the proof is very similar to that of [2, Theorem 5.1].
So, it is omitted. h

Proofs of Propositions 4.1,4.2. Clearly, Propositions 4.1,4.2 follow from Lemmas 4.3, 4.4, 4.6 and 4.9.

5. Discussion

In recent years, several authors investigated single-species population reaction–diffusion models where only part of the
population is migrating and another part is stationary. In this paper, following [5,8,18,19,21], a delayed reaction–diffusion
system with a quiescent stage is studied.
The phenomenon of biological invasion is investigated using traveling wave theory. We first transform the corresponding
wave system into a scalar delayed differential equation with an integral term. Then we establish the existence of traveling
wave fronts with speed c if c P cmin. Such an existence result implies that there is a transition zone moving from the trivial
steady state with no individuals to the coexistence steady state with both individuals of mobile and stationary classes, which
means that the invasion of both mobile and stationary classes can be successful. Furthermore, we establish the non-
existence, asymptotic behavior, monotonicity and uniqueness of traveling wave solutions. In particular, the effects of the
delay s and transfer rates c1, c2 on cmin are studied and we obtain some interesting results.
It should be mentioned that the monotone assumption @ 2f(u, v) P 0 for (u, v) 2 [0, K]2 is a crucial one in our main results.
However, the reproduction function in a model system arising from a practical problem may not satisfy the condition. For
example, f(u(x, t),u(x, t  s)) = du(x, t) + pu(x, t)eau(x,ts) for p/d > e, and
 
uðx; t  sÞ
f ðuðx; tÞ; uðx; t  sÞÞ ¼ uðx; tÞ 1  ;
K
do not satisfy the condition. When this monotone condition does not hold, the problem on traveling waves becomes much
harder due to lack of quasi-monotonicity. In this case, the existence of traveling waves for small values of the delay could be
obtained by using the idea of the so-called exponential ordering for delayed differential equations as in Huang and Zou [6]
and Wu and Zou [12,13]. However, the monotonicity and uniqueness of traveling waves of such non-monotone delayed
reaction–diffusion systems seem to be very interesting and challenging problems, see e.g., Ma and Zou [11] and the surveys
of Gourley et al. [3,4].
A problem close to traveling wave solutions is the so-called spread rate. The spread rate is a number c⁄ such that, for ini-
tial data with compact support, an observer traveling with speed c > c⁄ will eventually see the trivial state, and an observer
traveling with speed 0 < c < c⁄ will see the non-trivial state. For a large class of evolution systems (under a subtangential con-
dition), it has been shown that the spread rate equals the minimal speed of traveling waves. We expect that, for the delayed
reaction–diffusion system (1.2) with a quiescent stage, the spread rate c⁄ coincides with the minimal wave speed cmin. The
work due to [14] maybe be a good reference for this problem. We leave these questions for future research.

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