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CMS Books in Mathematics
Canadian
Mathematical Society
Société mathématique
Dynamical Systems
in Population
Biology
Second Edition
Canadian Mathematical Society
Société mathématique du Canada
Editors-in-Chief
Rédacteurs-en-chef
K. Dilcher
K. Taylor
Advisory Board
Comité consultatif
M. Barlow
H. Bauschke
L. Edelstein-Keshet
N. Kamran
M. Kotchetov
Dynamical Systems
in Population Biology
Second Edition
123
Xiao-Qiang Zhao
Department of Mathematics and Statistics
Memorial University of Newfoundland
St. John’s, NL, Canada
Mathematics Subject Classification (2010): 34Cxx, 34Kxx, 35Bxx, 35R10, 37Bxx, 37Cxx, 37N25,
39A05, 92D25, 92D30, 92D40
du1
= f1 (t, u1 , u2 ),
dt (0.1)
du2
= f2 (t, u1 , u2 ),
dt
where f1 and f2 are continuously differentiable and ω-periodic in t, and
∂fi /∂uj ≤ 0, i = j. We assume that, for each v ∈ R2 , the unique solution
u(t, v) of system (0.1) satisfying u(0) = v exists globally on [0, ∞).
vii
viii Preface
We use the proof provided in [334, Theorem 7.4.2]. Indeed, it suffices to prove
that every bounded orbit of {P n } converges to a fixed point of P . Given two
points u, v ∈ R2 , one or more of the four relations u ≤ v, v ≤ u, u ≤K v,
v ≤K u must hold. Now, if P n0 u0 ≤K P n0 +1 u0 (or the reverse inequality)
holds for some n0 ≥ 0, then (P1) implies that P n u0 ≤K P n+1 u0 (or the
reverse inequality) holds for all n ≥ n0 . Therefore, {P n u0 } converges to some
fixed point ū, since the sequence is bounded and eventually monotone. The
proof is complete in this case, so we assume that there does not exist such an
n0 as just described. In particular, it follows that u0 is not a fixed point of
P . Then it follows that for each n we must have either P n+1 u0 ≤ P n u0 or
the reverse inequality. Suppose for definiteness that u0 ≤ P u0 , the other case
being similar. We claim that P n u0 ≤ P n+1 u0 for all n. If not, there exists n0
such that
u0 ≤ P u0 ≤ P 2 u0 ≤ · · · ≤ P n0 −1 u0 ≤ P n0 u0
but P n0 u0 ≥ P n0 +1 u0 . Clearly, n0 ≥ 1 since u0 ≤ P u0 . Applying (P2) to the
displayed inequality yields P n0 −1 u0 ≥ P n0 u0 and therefore P n0 −1 u0 = P n0 u0 .
Since P is one to one, u0 must be a fixed point, in contradiction to our
assumption. This proves the claim and implies that the sequence {P n u0 }
converges to some fixed point ū.
Preface ix
It is hoped that the reader will appreciate the elegance and simplicity of
the arguments supporting the above theorem, which are motivated by a now
classical paper of deMottoni and Schiaffino [92] for the special case of periodic
Lotka–Volterra systems. This example also illustrates the roles that Poincaré
maps and monotone discrete dynamical systems may play in the study of pe-
riodic systems. For certain nonautonomous perturbations of a periodic system
(e.g., an asymptotically periodic system), one may expect that the Poincaré
map associated with the unperturbed periodic system (e.g., the limiting pe-
riodic system) should be very helpful in understanding the dynamics of the
original system. For a nonperiodic nonautonomous system (e.g., almost peri-
odic system), we are not able to define a continuous or discrete-time dynamical
system on its state space. The skew-product semiflow approach has proved to
be very powerful in obtaining dynamics for certain types of nonautonomous
systems (see, e.g., [303, 300, 311]).
The main purpose of this book is to provide an introduction to the the-
ory of periodic semiflows on metric spaces and its applications to population
dynamics. Naturally, the selection of the material is highly subjective and
largely influenced by my personal interests. In fact, the contents of this book
are predominantly from my own and my collaborators’ recent works. Also, the
list of references is by no means exhaustive, and I apologize for the exclusion
of many other related works.
Chapter 1 is devoted to abstract discrete dynamical systems on metric
spaces. We study global attractors, chain transitivity, strong repellers, and
perturbations. In particular, we will show that a dissipative, uniformly persis-
tent, and asymptotically compact system must admit a coexistence state. This
result is very useful in proving the existence of (all or partial componentwise)
positive periodic solutions of periodic evolutionary systems.
The focus of Chapter 2 is on global dynamics in certain types of monotone
discrete dynamical systems on ordered Banach spaces. Here we are interested
in the abstract results on attracting order intervals, global attractivity, and
global convergence, which may be easily applied to various population models.
In Chapter 3, we introduce the concept of periodic semiflows and prove
a theorem on the reduction of uniform persistence to that of the associated
Poincaré map. The asymptotically periodic semiflows, nonautonomous semi-
flows, skew-product semiflows, and continuous processes are also discussed.
In Chapter 4, as a first application of the previous abstract results, we
analyze in detail a discrete-time, size-structured chemostat model that is de-
scribed by a system of difference equations, although in this book our main
concern is with global dynamics in periodic and almost periodic systems. The
reason for this choice is that we want to show how the theory of discrete dy-
namical systems can be applied to discrete-time models governed by difference
equations (or maps).
In the rest of the book, we apply the results in Chapters 1–3 to continuous-
time periodic population models: in Chapter 5 to the N -species competition
in a periodic chemostat, in Chapter 6 to almost periodic competitive systems,
x Preface
For this edition, I have corrected some typos, revised Sections 1.1 and 1.3.4 by
using the concepts of global attractors and strong global attractors, deleted
the original subsection on order persistence from Section 1.3, and added three
new sections about persistence and attractors (Section 1.3.3), saddle point
behavior for monotone semiflows (Section 2.5), and solution maps of abstract
nonautonomous functional differential equations (Section 3.5), respectively. I
have also mentioned more related references in the notes sections of Chap-
ters 1–10.
In addition, I have added four new chapters. Chapter 11 is devoted to the
general theory of basic reproduction ratios R0 for compartmental models of
periodic functional differential equations and autonomous reaction–diffusion
systems. Chapter 12 deals with the threshold dynamics in terms of R0 for a
new class of population models with time periodic delays. In Chapter 13, we
study a periodic reaction–diffusion SIS system and investigate the effect of
spatial and temporal heterogeneities on the extinction and persistence of the
infectious disease. The final chapter, Chapter 14, provides a complete analysis
of the disease-free dynamics and global dynamics for a nonlocal spatial model
of Lyme disease. It is expected that Chapters 12–14 may serve as templates for
future investigations on other population models with spatial and temporal
heterogeneities.
My sincere thanks goes to Jifa Jiang, Xing Liang, Yijun Lou, Pierre Magal,
Rui Peng, Wendi Wang, and Xiao Yu, whose joint research articles with me
have been incorporated in the second edition. I am also very grateful to all
collaborators and friends for their encouragements, suggestions, and assistance
with this revision.
xi
Contents
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
2 Monotone Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
2.1 Attracting Order Intervals and Connecting Orbits . . . . . . . . . . . 44
2.2 Global Attractivity and Convergence . . . . . . . . . . . . . . . . . . . . . . . 48
2.3 Subhomogeneous Maps and Skew-Product Semiflows . . . . . . . . . 52
2.4 Competitive Systems on Ordered Banach Spaces . . . . . . . . . . . . 59
2.5 Saddle Point Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
2.6 Exponential Ordering Induced Monotonicity . . . . . . . . . . . . . . . . 69
2.7 Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
3 Nonautonomous Semiflows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
3.1 Periodic Semiflows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
3.1.1 Reduction to Poincaré Maps . . . . . . . . . . . . . . . . . . . . . . . . 78
3.1.2 Monotone Periodic Systems . . . . . . . . . . . . . . . . . . . . . . . . . 80
xiii
xiv Contents
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
1
Dissipative Dynamical Systems
So the family {An }n≥0 satisfies the conditions of assertion (a) in Lemma 1.1.2,
and we deduce that ω(B) is nonempty, compact, and δ(An , ω(B)) → 0, as
n → +∞. So ω(B) attracts B for f . Moreover, we have
⎛ ⎞
f⎝ f m (B)⎠ = f m (B), ∀n ≥ 0,
m≥n m≥n+1
for any nonempty closed bounded subset B ⊂ X for which f (B) ⊂ B. This
implies that any α-contracting map is asymptotically smooth.
A positively invariant subset B ⊂ X for f is said to be stable if for any
neighborhood V of B, there exists a neighborhood U ⊂ V of B such that
f n (U ) ⊂ V, ∀n ≥ 0. We say that A is globally asymptotically stable for f if,
in addition, A attracts points of X for f .
By the proof that (i) implies (ii) in [141, Theorem 2.2.5], we have the
following result.
Lemma 1.1.4. Let B ⊂ X be compact, and positively invariant for f . If B
attracts compact subsets of some neighborhood of itself, then B is stable.
Definition 1.1.3. A nonempty, compact, and invariant set A ⊂ X is said to
be an attractor for f if A attracts some open neighborhood of itself; a global
attractor for f if A is an attractor that attracts every point in X; and a strong
global attractor for f if A attracts every bounded subset of X.
Remark 1.1.1. The notion of attractor and global attractor was used in
[164, 304]. The strong global attractor was defined as global attractor in
[141, 358, 286]. In the case where the dimension of X is finite, it is easy to see
that both global attractor and strong global attractor are equivalent. In the
infinite-dimensional case of X, however, there exist discrete- and continuous-
time dynamical systems that admit global attractors, but no strong global
attractors, see Example 1.3.3 and [241, Sections 5.1–5.3].
The following result is essentially the same as [142, Theorem 3.2]. Note
that the proof of this result was not provided in [142]. For completeness, we
state it in terms of global attractors and give an elementary proof below.
Theorem 1.1.2. (Global Attractors) Let f : X → X be a continuous
map. Assume that
(a) f is point dissipative and asymptotically smooth;
(b) Positive orbits of compact subsets of X for f are bounded.
Then f has a global attractor A ⊂ X. Moreover, if a subset B of X admits
the property that γ + (f k (B)) is bounded for some k ≥ 0, then A attracts B
for f .
Proof. Assume that (a) is satisfied. Since f is point dissipative, we can find
a closed and bounded subset B0 in X such that for each x ∈ X, there exists
k = k(x) ≥ 0, f n (x) ∈ B0 , ∀n ≥ k. Define
J(B0 ) := {y ∈ B0 : f n (y) ∈ B0 , ∀n ≥ 0} .
Thus, f (J(B0 )) ⊂ J(B0 ), and for every x ∈ X, there exists k = k(x) ≥ 0 such
that f k (x) ∈ J(B0 ). Since J(B0 ) is closed and bounded, and f is asymptot-
ically smooth, Lemma 1.1.3 implies that ω(J(B0 )) is compact invariant, and
attracts points of X.
6 1 Dissipative Dynamical Systems
Assume, in addition, that (b) is satisfied. We claim that there exists an ε >
0 such that γ + (N (ω(J(B0 )), ε)) is bounded. Assume, by contradiction, that
1
γ + N ω(J(B0 )), n+1 is unbounded for each n > 0. Let z ∈ X be fixed.
1
Then we can find a sequence xn ∈ N ω(J(B0 )), n+1 , and a sequence of
integers mn ≥ 0 such that d(z, f mn
(xn )) ≥ n. Since ω(J(B0 )) is compact,
we can always assume that xn → x ∈ ω(J(B)), as n → +∞. Since H :=
{xn : n ≥ 0}∪{x} is compact, assumption (b) implies that γ + (H) is bounded,
a contradiction. Let D = γ + (N (ω(J(B0 )), ε)). Then D is closed, bounded,
and positively invariant for f . Since ω(J(B0 )) attracts points of X for f , and
ω(J(B0 )) ⊂ N (ω(J(B0 )), ε) ⊂ int(D), we deduce that for each x ∈ X, there
exists k = k(x) ≥ 0 such that f k (x) ∈ int(D). It then follows that for each
compact subset C of X, there exists an integer k ≥ 0 such that f k (C) ⊂ D.
Thus, the set A := ω(D) attracts every compact subset of X. Fix a bounded
neighborhood V of A. By Lemma 1.1.4, it follows that A is stable, and hence,
there is a neighborhood W of A such that f n (W ) ⊂ V, ∀n ≥ 0. Clearly, the
set U := ∪n≥0 f n (W ) is a bounded neighborhood of A, and f (U ) ⊂ U . Since
f is asymptotically smooth, there is a compact set J ⊂ U such that J attracts
U . By Lemma 1.1.3, ω(U ) is nonempty, compact, invariant for f , and attracts
U . Since A attracts ω(U ), we have ω(U ) ⊂ A. Thus, A is a global attractor
for f .
To prove the last part of the theorem, without loss of generality we assume
that B is a bounded subset of X and γ + (B) is bounded. We set K = γ + (B).
Then f (K) ⊂ K. Since K is bounded and f is asymptotically smooth, there
exists a compact C which attracts K for f . Note that f k (B) ⊂ f k (γ + (B)) ⊂
f k (K) , ∀k ≥ 0. Thus, C attracts B for f . By Lemma 1.1.3, we deduce that
ω(B) is nonempty, compact, invariant for f and attracts B. Since A is a
global attractor for f , it follows that A attracts compact subsets of X. By the
invariance of ω(B) for f , we deduce that ω(B) ⊂ A, and hence, A attracts B
for f .
Remark 1.1.2. From the first part of the proof of Theorem 1.1.2, it is easy to
see that if f is point dissipative and asymptotically smooth, then there exists
a nonempty, compact, and invariant subset C of X for f such that C attracts
every point in X for f .
The following lemma provides sufficient conditions for the positive orbit
of a compact set to be bounded.
Proof. Since f is point dissipative, we can choose a bounded and open subset
V of X such that for each x ∈ X there exists n0 = n0 (x) ≥ 0 such that
1.1 Limit Sets and Global Attractors 7
which is impossible. In the case where the sequence {lp } is unbounded, there
exists a subsequence lpk → ∞ as k → ∞. Then for each fixed m ≥ 1, there
exists an integer km such that m ≤ lpk , ∀k ≥ km , and hence,
f m (ypk ) ∈ X \ V, ∀k ≥ km .
Letting k → ∞, we obtain
f m (y) ∈ X \ V, ∀m ≥ 1,
Remark 1.1.3. It is easy to see that a metric space (X, d) is complete if and
only if for any subset B of X, α(B) = 0 implies that B is compact. How-
ever, we can prove that Lemmas 1.1.3 and 1.1.4 also hold for non-complete
metric spaces by employing the equivalence between the compactness and the
sequential compactness for metric spaces. It then follows that Theorems 1.1.2
and 1.1.3 are still valid for any metric space. We refer to [64, 286] for the
existence of strong global attractors of continuous-time semiflows on a metric
space.
Tn = Sn−1 ◦ Sn−2 ◦ · · · ◦ S1 ◦ S0 , n ≥ 1.
The orbit of x ∈ X under this process is the set γ + (x) = {Tn (x) : n ≥ 0},
and its omega limit set is
ω(x) = y ∈ X : ∃nk → ∞ such that lim Tnk (x) = y .
k→∞
xn+1 = f (n, xn ), n ≥ 0,
T0 = I, Tn = Sn−1 ◦ · · · ◦ S1 ◦ S0 : X → X, n ≥ 1,
Lemma 1.2.3. The omega limit set of any precompact asymptotic pseudo-
orbit of a continuous map S : X → X is nonempty, compact, invariant, and
internally chain transitive.
Proof. Let (N+ , ρ) be the compact metric space defined in the proof of
Lemma 1.2.2. Let {xn : n ≥ 0} be a precompact asymptotic pseudo-orbit
of S : X → X, and denote its compact omega limit set by ω. Define a metric
space
Y = ({∞} × X) ∪ {(n, xn ) : n ≥ 0}
12 1 Dissipative Dynamical Systems
and
By Definition 1.2.3 and the fact that d(xn+1 , S(x)) ≤ d(xn+1 , S(xn )) +
d(S(xn ), S(x)) for x ∈ X, n ≥ 0, it easily follows that g : Y → Y is con-
tinuous. Let γ + (0, x0 ) = {(n, xn ) : n ≥ 0} be the positive orbit of (0, x0 ) for
the discrete semiflow g n : Y → Y, n ≥ 0. Then γ + (0, x0 ) is precompact in Y ,
and its omega limit ω(0, x0 ) is {∞} × ω, which by Lemma 1.2.1 is invariant
and internally chain transitive for g. Applying the definition of g, we see that
ω is invariant and internally chain transitive for S.
Let A and B be two nonempty compact subsets of X. Recall that the
Hausdorff distance between A and B is defined by
d(Sn (u), S(v)) ≤ d(Sn (u), S(u)) + d(S(u), S(v)) < /3.
Fix n > N such that dH (Dn , D) < δ. For any a, b ∈ D, there are points
x, y ∈ Dn such that d(x, a) < δ and d(y, b) < δ. Since Dn is internally chain
transitive for Sn , there is a δ-chain {z1 = x, z2 , . . . , zm+1 = y} in Dn for Sn
connecting x to y. For each i = 2, . . . , m we can find wi ∈ D with d(wi , zi ) < δ,
since Dn is contained in the δ-neighborhood of D. Let w1 = a, wm+1 = b. We
then have
Proof. Let ω = ω(x) be the omega limit set of a precompact orbit γ(x) =
{Φ(t)x : t ≥ 0} in X. Then ω is nonempty, compact, invariant and
limt→∞ d(Φ(t)x, ω) = 0. Let > 0 and t0 > 0 be given. By the uniform
continuity of Φ(t)x for (t, x) in the compact set [t0 , 2t0 ] × ω, there is a
δ = δ(, t0 ) ∈ (0, 3 ) such that for any t ∈ [t0 , 2t0 ] and u and v in the open
δ-neighborhood U of ω with d(u, v) < δ, we have d(Φ(t)u, Φ(t)v) < 3 . It
then follows that there exists a sufficiently large T0 = T0 (δ) > 0 such that
Φ(t)x ∈ U , for all t ≥ T0 . For any a, b ∈ ω, there exist T1 > T0 and T2 > T0
with T2 > T1 + t0 such that d(Φ(T1 )x, Φ(t0 )a) < 3 and d(Φ(T2 )x, b) < 3 . Let
m be the greatest integer that is not greater than T2t−T 0
1
. Then m ≥ 1. Set
and
ti = t0 for i = 1, . . . , m; tm+1 = T2 − T1 − (m − 1)t0 .
Then tm+1 ∈ [t0 , 2t0 ). It follows that d(Φ(ti )yi , yi+1 ) <
3 for all i = 1, . . . , m+
1. Thus the sequence
With Lemma 1.2.1 it is easy to see that there are analogues of Lem-
mas 1.2.2 and 1.2.3 for continuous-time semiflows. The following result is an
analogue of Lemma 1.2.4 for continuous-time semiflows.
d(Φti (wi ), wi+1 ) ≤ d(Φti (wi ), Φnti (zi )) + d(Φnti (zi ), zi+1 ) + d(zi+1 , wi+1 )
< /3 + δ + δ <
Example 1.2.2. Note that if in Lemma 1.2.4 Dn is an omega limit set for
Φn (and therefore internally chain transitive by Lemma 1.2.1 ), the set D
need not be an omega limit set for the limit semiflow Φ, although it must be
chain transitive. Easy examples are constructed with Φn = Φ, ∀n ≥ 1. For
example, consider the flow generated by the planar vector field given in polar
coordinates by
r = 0, θ = 1 − r.
The unit circle D = {r = 1}, consisting of equilibria, is chain transitive but is
not an omega limit set of any point, yet D is the Hausdorff limit of the omega
limit sets Dn = {r = 1 + n1 }.
Proof. The sufficiency follows from Lemma 1.2.3. To prove the necessity,
we can choose a point x ∈ M since M is nonempty. For any > 0,
the compactness of M implies that there is a finite sequence of points
1.2 Chain Transitivity and Attractivity 15
Proof. Since M is an isolated invariant set, there exists an > 0 such that
M is the maximal invariant set in the closed -neighborhood of M . By the
assumption, we can choose a ∈ L ∩ M and b ∈ L with d(b, M ) > . For
any integer k ≥ 1, by the internal chain transitivity of L, there exists a
1 1
k -chain {y1 = a, . . . , ylk +1 = b} in L connecting a and b, and a k -chain
k k
by