Uniform persistence and permanence for non-autonomous
semi¯ows in population biology
qHorst R. Thieme
*Department of Mathematics, Arizona State University, Tempe, AZ 85287-1804, USA
Received 21 February 2000; accepted 19 April 2000
Abstract
Conditions are presented for uniform strong persistence of non-autonomous semi¯ows, taking uniform weak persistence for granted. Turning the idea of persistence upside down, conditions are derived for non-autonomous semi¯ows to be point-dissipative. These results are applied to time-heterogeneous models of S±I±R±S type for the spread of infectious childhood diseases. If some of the parameter functions are as-ymptotically almost periodic, an almost sharp threshold result is obtained for uniform strong endemicity versus extinction in terms of asymptotic time averages. Applications are also presented to scalar retarded functional dierential equations modeling one species population growth. Ó 2000 Elsevier Science Inc. All rights reserved.
MSC:34C35; 34D05; 34D40; 34K25; 92D25; 92D30
Keywords: Persistence; Permanence; Dissipativity; Dynamical systems; Epidemic models; Functional dierential equations; (Asymptotically) almost periodic functions; Time averages
1. Introduction
Persistence (or permanence) is an important property of dynamical systems and of the systems in ecology, epidemics etc., they are modeling. Persistence addresses the long-term survival of some or all components of a system, while permanence also deals with the limits of growth for some (or all) components of the system, For background information and refer-ences we refer to Thieme [40]. We show that uniform weak persistence implies uniform (strong)
q
Research partially supported by NSF grants DMS-9403884 and DMS-9706787.
*Tel.: +1-480 965 4772; fax: +1-480 965 8119. E-mail address:h.thieme@asu.edu (H.R. Thieme).
0025-5564/00/$ - see front matter Ó 2000 Elsevier Science Inc. All rights reserved.
persistence. Loosely speaking, a population is uniformly weakly persistent if its size, while it may come arbitrarily close to 0 every now and then, always climbs back to a level that eventually is independent of the initial data. The population is uniformly (strongly) persistent, if its size is bounded away from 0 and the bound does not depend on the initial data after suciently long time. The population is permanent, if it is uniformly (strongly) persistent and if the population size is bounded with the bound not depending on the initial data after su-ciently long time.
While a general persistence/permanence theory is available for autonomous semi¯ows [17,23] and special non-autonomous systems have been considered in the past [3,41] (for more refer-ences see [40]), a general non-autonomous theory is still under development. The importance of a non-autonomous theory is obvious because all ®eld populations live in a seasonal environ-ment. The approach in this paper is based on uniform weak persistence [11,39]; alternatively one can try to reduce the non-autonomous case to the autonomous one by using skew product ¯ows [43]. Both approaches show the existence of positive lower bounds which, for suciently large times, do not depend on the initial conditions, but they do not provide estimates of these bounds.
It is worth mentioning that permanence of the biological system in particular involves the point-dissipativity of the semi¯ow (existence of a bounded absorbing set) that models the dy-namics of the system. While point-dissipativity has often been assumed to prove persistence [17], we will in turn use persistence techniques to derive conditions for point-dissipativity.
The results for non-autonomous semi¯ows cannot be so elegantly stated as in the autonomous case [11,39], as we need additional conditions which appear quite technical though they can ef-fectively be checked in many applications. As a trade-o, compactness requirements for the state space or at least for an attracting set can be replaced by appropriate equi-continuity conditions for the semi¯ow.
We apply our results to establish threshold criteria for disease extinction and disease persistence in time-hetergeneous S±I±R±S epidemic models and to establish permanence for a one species model consisting of a scalar retarded functional dierential equation.
This paper is organized as follows. In Section 2, we generalize the result in [40] that uniform weak persistence implies uniform strong persistence under appropriate extra conditions. Among other things, it now covers situations with relaxed invariance (cf. [10, Section 6]). We show how persistence theory can be turned around to show ultimate boundedness (or point dissipativity) for non-autonomous semi¯ows. We also illustrate the versatility of the framework by establishing persistence that holds uniformly with respect to parameters. This is related to robust permanence [21,32], where permanence of semi¯ows induced by ordinary dierential equations is preserved
under small Cr perturbations of the vector ®eld.
In Section 3, we derive threshold results for disease extinction and disease persistence for an unstructured epidemic model of S±I±R±S type. In Section 4, we show that these results also hold if the sojourn time in the removed class has a general distribution. In Section 5, we study a time-heterogeneous model for the dynamics of one species with general feed-back, formulated by a scalar retarded functional dierential equation
_
We derive conditions for permanence, i.e., for the existence of constants 0< <c<1such that
6lim inf
t!1 x t6lim supt!1 x t6c
with;c not depending on the initial conditionsxr / as long asx r>0.
The conditions we obtain involve asymptotic time averages and are reminiscent of conditions obtained by Burton and Hutson [3] for prey±predator models and by Wu et al. [41] for almost periodic Kolmogorov equations. We have collected some material on asymptotic time averages and their connection to (asymptotic) almost periodicity in Appendix A. Each application section (Sections 3±5) presents a typical persistence or permanence result shortly after the explanation of the model equations. These results are not the most general possible, but their meaning can be grasped independently of Section 2 and the rest of the respective sections.
2. Uniform weak is uniform strong
LetX be a set,r0 2R, and
D f t;s;r06s6t<1g:
A mappingW:DX !X is called a (non-autonomous) semi¯owon X(anchored atr0) if
W t;s;W s;r;x W t;r;x; W r;r;x x 8tPsPrPr0; x2X:
IfXis topological space and the mappingWis continuous and a semi¯ow,Wis called acontinuous
semi¯ow.The semi¯ows we will consider are not necessarily continuous.
Wis called an autonomous semi¯ow ifW tr;r;xdoes not depend onrPr0fortP0; x2X.
Further let
q:X ! 0;1
be a non-negative functional onXand
XqX \ fq>0g:
Xq is not necessarily forward invariant under W. We consider the function
r:0;1 X r0;1 ! 0;1
de®ned by
r t;x;r q W tr;r;x; tP0; x2X; rPr0: 2:1
We make the following assumption throughout this section, namely that the real-valued function
r ;x;r is continuous on0;1for all x2X; rPr0.
We notice the following relation between rand W:
r t;W s;r;x;s q W ts;s;W s;r;x q W ts;r;x;
r 0;x;r q x: 2:2
We introduce the following notation:
r1 x;r lim sup
t!1
r t;x;r; r1 x;r lim inf
De®nition 2.1. Wis called
· weaklyq-persistentif
r1 x;r>0 8x2Xq; rPr0;
· strongly q-persistentif
r1 x;r>0 8x2Xq; rPr0;
· uniformly weakly q-persistentif there exists some >0 such that
r1 x;r> 8x2Xq; r r0;
· uniformly (strongly) q-persistentif there exists some >0 such that
r1 x;r> 8x2Xq; r r0;
· weaklyq-dissipative if there exists somec>0 such that
r1 x;r<c 8x2X; rPr0;
· strongly q-dissipativeif there exists some c>0 such that
r1 x;r<c 8x2X; rPr0;
· q-permanent if Wis both uniformly stronglyq-persistent and stronglyq-dissipative.
If no misunderstanding about the functional q is possible, we use persistent rather than
q-persistent etc.
In the topologically oriented persistence theory, q x is the distance of the point x from a
certain set, the boundary of extinction. Freeingqfrom this interpretation makes it possible to use
persistence techniques to derive strong from weak dissipativity. It will also come in handy for studying persistence in physiologically structured population models, where the appropriate state spaces are formed by regular Borel measures on a locally compact Hausdor space and the
semi¯ows are continuous not in the strong, but in the weak topology (see [5] for the linear
foundation of the construction of such semi¯ows and [6] for a ®rst non-linear extension). Obviously uniform strong persistence implies uniform weak persistence. Deriving conditions for the converse to hold is not only of theoretical but also of practical interest, because uniform weak persistence can often be checked in concrete situations with relative ease. Here the following remark is helpful.
Remark.Wis uniformly weaklyq-persistent if and only if, for suciently small >0, no x2Xq
and no rPr0 can be found such that r t;x;r6 for all tP0.
also [12,42]). Actually one can replace compactness completely by equi-continuity requirements
forr. This makes it possible to use persistence techniques to derive conditions for
point-dissip-ativity of semi¯ows.
We will still use the idea of an absorbing or attracting set, but we will not assume that this set is bounded or even compact.
LetBbe a subset ofX;J Nor an interval of the forma;1. We will associate two dierent
interpretations with the notation
xj!B as j! 1;
where xjj2J is a sequence or family of elements inX. The ®rst interpretation is often appropriate
for semi¯ows that operate on sets rather than topological spaces, or on locally compact spaces, or
come from problems with ®nite memory. We say thatB absorbs xj, if
/ there exists somej0 2J such that xj2Bfor all j2J; jPj0.
For semi¯ows on topological spaces that come from problems with in®nite memory (e.g., age-dependent equations without ®nite maximum age), the following interpretation is more
appro-priate: We say thatB attracts xj, if
. for any open setU B there exists some jU 2J such that xj2U for all j2J; jPjU.
IfX is a metric space,B attracts xj if the distance ofxj fromB tends to 0 asj! 1.
Wis said to have property (CA) if the following holds:
(CA) There exists a subsetB inXwith the following two properties:
· For allx2Xq; sPr0, we haveW t;s;x !B; t! 1.
· If rj is a sequence of real numbers and yj a sequence inXsuch that rj! 1 andyj!B as
j! 1 and, for some >0; q yj for allj2N, then the continuity ofr ;yj;rjis uniform
inj2N, possibly after choosing a sub-sequence.
In both conditions, of course, `!B' has to be consistently interpreted as B being either
ab-sorbing or attracting. IfW t;s;x !B is interpreted in the ®rst way /, we say that Bis an
ab-sorbing setforW, while in the second interpretation . we say thatBis anattracting set.IfBis
absorbing, the second property of (CA) is more succinctly stated as the continuity ofr ;y;rto be
uniform inrPr0 andy 2B; q y for ®xed, but arbitrary >0.
Recall that we assume throughout this section thatr ;x;ris continuous for allx2X; rPr0.
The notation (CA) has been chosen to recall the historical connection to the existence of a compact attracting set. The following generalization has been proved useful in studying the
persistence of host and/or parasite populations in time-autonomous epidemic models [39]. IfXis
topological space, we call a subsetBofX relativelyq-compactif all its intersections with q-shells
have compact closure inX. Aq-shell is a set fx2X;16q x62gwith 0< 1 < 2<1.
Remark.(CA) holds if the following holds:
· W t;s;x W tÿs;0;x U tÿs;xis a continuous autonomous semi¯ow on a metric spaceX.
· There exists a relatively q-compact setBwithU t;x !B ast! 1 for all x2X.
· qis continuous onX ifBis absorbing, and uniformly continuous if Bis attracting.
Proof.Assume that r t;yj;sj q U t;yjis not continuous in t uniformly in yj with yj!B as
j! 1; q yj . After choosing a sub-sequence of yj we ®ndtP0 andtj!tasj! 1 such
d yj;zj !0 as j! 1. Sinceqis (uniformly) continuous on the metric spaceXandq yj for
large enough j, the zj are in the intersection of B with a q-shell and so have a converging
sub-sequence. Letting y be the limit and using the continuity of q and U, we have
q U tj;yj !q U t;y; q U t;yj !q U t;ycontradicting our previous statement.
Even in a locally compact metric space, uniform weak persistence will not imply uniform strong persistence if the semi¯ow is non-autonomous.
Example 2.2.LetX 0;1andq x x. Construct a pair of functionsx},aon0;1with the following properties:
_
x} ÿx}a t; tP0; x} 0 2;
0<x} t62; 06a t63 8tP0;
lim sup
t!1
x} t 2; lim inf
t!1 x} t 0:
Now let Wbe the semi¯ow induced by the solutions of
_
x ÿxa t; tPr; x r x0;
i.e., W t;r;x0 x t. Since x t ÿx} t !0 as t! 1, we have that Wis uniformly weakly
per-sistent, but not strongly perper-sistent, though Wis point-dissipative and asymptotically smooth if
considered on0;1.
This example teaches us that we need some condition that takes care of the non-autonomous nature of the semi¯ow.
For every >0; t>0 we de®ne setR andR t; as follows (Bis the absorbing or attracting
set in (CA)):
R t; consists of continuous functions r~:0;t ! 0; ;
~
r t 0<r~ 0 ;
~
r s lim
j!1r s;yj;sj uniformly in s2 0;t
for sequences sj inr0;1; yj inX withsj! 1; yj !B;asj! 1:
R consists of continuous functions r~:0;1 ! 0; ;
0<r~ 0 ;
~
r s lim
j!1r s;yj;sj locally uniformly in sP0
for sequences sj inr0;1; yj inX withsj! 1; yj !B;asj! 1:
2:3
The semi¯ow Wis said to have property (PS) if the following holds:
Remark.(PS) automatically holds for a continuous autonomous semi¯ow Uwhich is uniformly
weaklyq-persistent, provided that there is a relativelyq-compact attracting setBforUsuch that
Xq\B is forward invariant underU and that q is uniformly continuous on the metric space X.
This statement holdsMutandis mutatisfor asymptotically autonomous semi¯ows [40].
Proof.Following the proof of the Remark subsequent (CA) we notice that the functionsr~inR or R t; , respectively have the form r~ s q U s;y for some y2B; q y . R t; is empty
becauseXq\Bis forward invariant.R ;follows from the uniform weak persistence ofUfor
suciently small >0.
Theorem 2.3. If a semiflowWis uniformly weakly q-persistent and has properties(CA) and(PS), then it is uniformly strongly persistent.
Proof.Assume thatWis uniformly weakly persistent. Then there exist >0 such that
r1 x;r> 8x2Xq; rPr0:
By (PS) we can arrange that the setsR and R t; are empty for allt>0 for this .
Step 1: Letx2Xq and rPr0 such that lim inft!1r t;x;r< 1 < .
Sinceq W ;r;xis continuous onr;1, there exist sequences skinr0;1and tkin 0;1
such that sk! 1 k ! 1and
q W sk;r;x ; q W tksk;r;x< 1 8k2N;
q W ssk;r;x6 whenever 06s6tk:
Step 2:Suppose thatWis not uniformly strongly persistent.
Then there exist sequences rj inr0;1and xj 2Xq such that
lim inf
t!1 q W t;rj;xj !0; j! 1:
Using step 1 and a diagonalization procedure we obtain sequences rjand sjinr0;1, and tj
in0;1such that sjPrjjfor all j2N, and
q W sj;rj;xj ;
q W tjsj;rj;xj !0; j! 1;
q W ssj;rj;xj6; 06s6tj:
Since, for every j2N;W t;rj;xj !B as t! 1 by the ®rst property in (CA), we can further
achieve that
W sj;rj;xj !B; j! 1:
SetyjW sj;rj;xj. Using the de®nition of rin (2.1), we have the following situation:
q yj ;
r tj;yj;sj !0; j! 1;
r s;yj;sj6; 06s6tj;
We claim: tj ! 1 asj! 1.
If not, after choosing sub-sequences,tj!t andr ;yj;sj is continuous uniformly inj. By the
second assumption of (CA), there exist somed>0 such that
r s;yj;sj6; 06s6td
for suciently large j. By the Arzela±Ascoli theorem, after choosing another sub-sequence, we
have
~
r s lim
j!1r s;yj;sj6; 06s6t;
with convergence holding uniformly for s2 0;td. Obviously r~ 0 >0. Again using that
r ;yj;sj is continuous uniformly inj,
0lim sup
j!1
r t;yj;sj r~ t:
Hence we have found an element in R t; which is a contradiction to (PS).
So,tj ! 1asj! 1. LettP0 be arbitrary. Then tj>t for suciently largej and
Pr s;yj;sj 8s2 0;t;
ifjis large enough. This implies thatr ;yj;sjis bounded on ®nite intervals in0;1uniformly in
j2N. The second assumption in (CA) implies thatr ;yj;sjis continuous on0;1uniformly in
j, possibly after choosing a sub-sequence. By the Arzela±Ascoli theorem, there exists a functionr~
on 0;1such that, after choosing another sub-sequence,
r t;yj;sj !r~ t; j! 1; locally uniformly in tP0
and
~
r 0 Pr~ t 8tP0:
Since R t; ;for all t>0,r~is strictly positive. So we have an element r~2R which again
contradicts (PS).
The ¯exibility ofq-persistence (compared with the topological concept of persistence) allows to
prove boundedness results.
For every j>0; t>0 we de®ne sets R~ j and R~ t;j as follows (B is the absorbing or
at-tracting set in (CA)):
~
R j consists of continuous functions r~:0;1 ! j;1;
~
r 0 j; r~ s lim
j!1r s;yj;sj locally uniformly in sP0
for sequences sj inr0;1; yj inX with sj! 1; yj !B asj! 1:
~
R t;j consists of continuous functions r~:0;t ! j;1; ~
r 0 j; lim
s!tÿr~ s 1; ~
r s lim
j!1r s;yj;sj uniformly in s2 0;t
Theorem 2.4. Let Wbe a semiflow that has the property (CA).Assume that, if jis chosen
suffi-ciently large, the setsR~ j andR~ t;jare empty for all tP0.
ThenWis stronglyq-dissipative whenever it is weakly q-dissipative.
Proof.Assume the Wis weaklyq-dissipative. Set
~
q x 1 1q x:
Then q~ x>0 for allx2X and Wis uniformly weakly q~-persistent. Moreover W satis®es (CA)
and (PS) withq~replacingq. It follows from Theorem 2.3 thatWis uniformly stronglyq~-persistent,
i.e., stronglyq-dissipative.
In some applications one would like persistence to hold uniformly with respect to a parameter (cf. [9, end of Section 4], [36, Section 3] and [37, Section 4]).
LetN be an index set and assume that we have a family of (non-autonomous) semi¯ows
Wn :DX !X; n2N:
Wn is said to have property CAN if the following holds:
CAN There exists a subsetBinX with the following two properties:
· For all x2X; sPr0; n2N, we haveWn t;s;x !B; t! 1.
· If rjis a sequence of real numbers and yj a sequence inX; nja sequence inN
such thatrj! 1andyj!Bast! 1and, for some >0; q yj for allj2N,
then the continuity of
t7!q Wnj tsj;sj;yj
on 0;1is uniform in j2N, possibly after choosing a sub-sequence.
Setr t;y;r q Un tr;r;xfory x;n. For everytP0; >0, setsR andR t; are now
de®ned in obvious analogy to (2.3) using sequences yi xj;nj with xj! B. We say that the
family Wn satis®es property PSN if the following holds:
PSN If >0 is chosen suciently small, the setsR and R t; are empty for alltP0.
Theorem 2.5. Let N be a set and Wn; n2N, be a family of semiflows on X that has properties
CAN and PSN with absorbing or attracting set B. Further assume that
t7!q Wn ts;s;x
is continuous on 0;1for any x2X; n2N; sPr0. Finally assume there exists some >0 such
that
lim sup
t!1
q Wn t;s;xP 8x2Xq; n2N; sPr0:
Then there exists some >0 such that
lim inf
Proof.LetZ X N and
~
q x;n q x:
De®ne a semi¯owW~ :DZ !Z by
~
W t;s; x;n Wn t;s;x;n:
Then W~ is uniformly weaklyq~-persistent. Further W~ satis®es (CA) with absorbing or attracting
set BN (in the second case endow N with the trivial topology). One easily checks that the
other assumptions of Theorem 2.3 are satis®ed as well. It follows that W~ is uniformly strongly
~
q-persistent. This implies the assertion of the theorem.
Similarly one can prove boundedness results for families of semi¯ows.
Theorem 2.3 has an interesting twist for continuous autonomous semi¯owsU, whereXqis not
forward invariant under all, but only under total orbits ofU. We recall that/:R!X is a total
orbit of U, ifU t/ s / ts for all tP0; s2R.
Theorem 2.6.LetUbe a continuous autonomous semiflow on a metric space X which has a compact
attracting set K, i.e., a compact set K such thatdist U t;x;K !0; t! 1. We further assume for
any total orbit /:R!X of Uwith relatively compact range:
· Ifs2Randq / s>0, thenq / t>0 for allt>s.
ThenUis uniformly strongly q-persistent whenever it is uniformly weaklyq-persistent.
Proof. We can assume that the compact attracting setK is invariant under U. Otherwise we
re-place it by the closure of the union of allx-limit sets ofU. The assumption (CA) is satis®ed as we
have shown in the remark following (CA).
Using the compactness ofKin the metric spaceX, for every elementr~inR or inR t; we
haver~ s q U s;yfor somey 2K. So uniform weak persistence implies thatR is empty for
suciently small >0. Sincey 2KandKis invariant,U s;ycan be extended to a total orbit ofU
with relatively compact range. This implies R t; ;, because any r~ in this set would satisfy
~
r 0>0r~ t, while, by; r~ 0>0 would imply r~ t>0.
Situations whereXq is not forward invariant, but q-positivity is preserved by total orbits, are
met in stage-structured population or epidemic models, see, [10, Section 6].
Remark.Property holds, e.g., if the following two conditions are satis®ed:
(i) If/is a total orbit ofUwith relatively compact range,s2R, andq / r 0 for allr6s,
thenq / t 0 for all tPs.
(ii) There exist somes>0; d>0 such that q U t;x>0 for all t2 s;sd; x2Xq.
Proof.Ifx2Xq, it follows from the semi¯ow property and repeated use of (ii) thatq U t;x>0 for allt2 ms;m sd; m2N, and soq U t;x>0 for alltPms, wheremis the ®rst natural
number withmd>s. Let now/be a total orbit ofU; s2Randq / s>0. By (i), there exists a
sequence sjin ÿ1;swithsj! ÿ1asj! 1 and q / sj>0. Choose somesj<sÿms. By
To interpret condition (ii) of the above Remark, let us assume thatqUis the infection rate in an epidemic model. If the infection rate is positive at the beginning, it can be 0 somewhat later because all infectious individuals have recovered in the meanwhile and the infected ones are still in the latency period. Eventually however, after an elapse of time which only depends on the length of the various periods of infection, the infection rate will be positive again. Condition (i) is related to the fact that infectious diseases cannot come out of nothing. If infectives have not been around in the past, then they will not be in the future (excluding, of course, that they are introduced from outside, or that viruses mutate and so become able to infect the host species under consideration).
3. The time-heterogeneous S±I±R±S model
As an application of the theory developed in Section 2, we consider a model for the spread of infectious childhood diseases. It has been argued that the school system induces a
time-hetero-geneity in the per capita/capita infection rate,a, because the chain of infections is interrupted or at
least weakened by the vacations and new individuals are recruited into a scene with higher in-fection risk at the beginning of each school year [8,31]. Here we consider a model without exposed
period; the total population, with size N, is divided into its susceptible, S, infective, I, and
re-covered, R, parts, and the contraction of the disease is modeled by the law of mass action
in-volving susceptibles and infectives
N SIR;
dI
dt ÿl tI a tSIÿc tI; 3:1
dR
dt ÿl tRc tI ÿn tR:
We assume that the disease causes no fatalities and that the population size N t is a given
function of time t, l t is the instantaneous per capita mortality rate and c t and n t are the
instantaneous per capita rates of leaving the infective stage or removed stage, respectively.
In the ®rst step, one can assume thatais a periodic function and thatN;l;c and nare constant.
This case (sometimes for the S±I±R, sometimes for the S±E±I±R model with exposed stage) was studied numerically by London and Yorke [25] and Dietz [7], formally by Grossman et al. [13,14], and analytically by Smith [34,35] and Schwartz and Smith [33]. It was shown that periodic
so-lutions exist whose periods are integer multiples of the period ofaand that co-existence of stable
periodic solutions with dierent periods is possible [33±35]. In the S±E±I±R model, periodic forcing of the infection rate can even lead to a sequence of period doubling sub-harmonic bi-furcations and ®nally chaos [1,2,26,29,30]. Whether the same holds for the S±I±R model, is not known to me; Theorem 3.2 below suggests that the trajectories cannot be completely wild (see [8,19] for more detailed reviews).
Throughout this section we will assume that N;l;a;c; and n are arbitrary non-negative,
assumptions we derive conditions for extinction and persistence of the disease. In case that
N;l;a and c (but not necessarily n) are almost periodic and we will obtain an almost sharp
threshold result.
Unfortunately our approach does not seem to extend to the S±E±I±R±S model.
To set the stage, we recall the following result, if N;l;a;c; and n are constant [18]. If
R0aN= lc61; thenI t !0; ast! 1 for every solution of (3.1). IfR0 >1, there exists
a unique endemic equilibrium which attracts all solutions with I 0>0, in particular
I t !I >0 as t! 1 with a uniquely determined I >0. While we cannot retain the global
asymptotic stability, we can preserve the threshold result concerning disease extinction versus endemicity using asymptotic time averages.
A function f :0;1 !R is called equi-mean-convergent (or to have uniformly convergent
means) if the (asymptotic) mean value
f lim
t!1
1
t
Z tr
r
f sds
exists, is independent ofrand the convergence is uniform inr. Almost periodic functions are
equi-mean-convergent, and so are asymptotically almost periodic functions and weakly asymptotically
almost periodic solutions (see Appendix A). If aN; c; lare equi-mean-convergent, we de®ne
R0 aN
lc:
Theorem 3.1. Let N be uniformly continuous on0;1andaN; landcbe equi-mean-convergent.
(a) If R0 <1, the disease dies out, i.e., for every solution of (3.1) we have I t;
R t !0 as t! 1.
(b)IfR0>1,the disease persists uniformly strongly in the population, in the sense that there
ex-ists some >0such thatI1lim inf t!1I t> for all solutions of(3.1)withI r>0for some
rP0.
The next result shows that strong persistence implies that the solutions cannot be arbitrarily wild.
Theorem 3.2. Let N be uniformly continuous on0;1andaN; l andcbe equi-mean-convergent
andR0>1.Then
1
t
Z t 0
a sS sds!lc aN
R0 ; t! 1
and
1
t
Z t 0
a s I s R sds! aN ÿ lc aN 1
ÿ 1
R0
Proof.From (3.4) we obtain
The last equality follows fromI being bounded away from 0 and bounded. This implies the ®rst
statement, the second is now obvious.
We will obtain Theorem 3.1 as a special case of extinction and persistence results which do not
assume equi-mean-convergence. To get started we notice that, for every solution of (3.1),I tis
non-negative for alltPrP0 where it is de®ned wheneverI r 0. The same property holds for
R. Adding the dierential equations forI and R it is easy to see that every solution satis®es
I t R t6 maxfI r R r;supNg; t>rP0:
A standard continuation argument now tells us that, given non-negative initial data at timerP0,
the solutions are de®ned for alltr.
Though this is not important for our mathematical considerations, epidemiologically one
would likeS NÿIÿRto be non-negative as well. This can be achieved by assuming thatNis
continuously dierentiable and that the population birth rate,B, satis®es
B t: d
dtN t l tN tP0: 3:2
Then SN ÿIÿRsatis®es the dierential equation
dS
dt B t ÿl tSÿa tSIn tR
from which we learn that, wheneverS rP0; rP0; thenS tP0 for alltPr. This implies
06S t;I t;R t6N t 8tPr:
Integrating theI-equation we obtain
1
Theorem 3.3. Let R} <1. Then the disease dies out, i.e., for every solution of (3.3) we have
I t; R t !0 ast! 1.
Proof. For simplicity we restrict the proof to a solution with initial data given at time 0. Since
S6N and R}<1, there exists some >0; r>0 such that
1
t
Z t
0
a sS sdsÿ1
t
Z t
0
c s l sds6 ÿ 8tPr:
By (3.3),
1
t ln I t
I r6 ÿ 8tPr:
Hence
I t6I reÿt; tPr
and I t !0; t! 1. The R-equation in (3.1) implies R t !0 as t! 1.
In order to obtain a weak persistence result we set
aN} lim inf
t!1
1
t
Z t 0
a sN sds;
c} lim sup
t!1
1
t
Z t 0
c sds; l} ;
R} aN}
l}c}
and let I1 be the limit superior ofI t ast! 1.
Theorem 3.4. LetR}>1.Then the disease uniformly weakly persists in the population, in the sense
that there exists some >0such thatI1> for all solutions of(3.1)withI r>0for somerP0.
does not depend on n.
Proof. Let us suppose that for every >0, there is some solution with I1< : Since R16 sup c=lI1;
I1R1<c
withc>0 being independent of the solution, of, and ofn. ByN SIR and (3.3),
d
for suciently larget: Hence there exists some t >0 such that
For suciently large timet, we have
1
In order to formulate a strong persistence results, letI1 be the limit inferior ofI t: Further
R: aN
Theorem 3.5. Let N be uniformly continuous on 0;1 and R>1: Then the disease uniformly
strongly persists in the population, in the sense that there exists some >0such that I1> for all
solutions of(3.1) withI r>0 for somerP0.LetI1 be the limit inferior of I t ast! 1.
endowed with the standard metric. Further we set
q I;R I:
In particular
W t;s;x I t;R t; tPsP0;
whereI;R solve (3.4) and I s;R s x. Further
r t;x;s I ts:
It may happen that, for certain initial data, we get solutions that are not epidemiologically
meaningful because I t R t>N t may occur, but this is irrelevant mathematically.
Condition (CA) is satis®ed because
I1<supN; R16supc
inflI
1
and the set
B fx I;R;I;R>0;I R6supNg
is absorbing and forward invariant. A standard Gronwall argument implies that r ;x;s is
continuous on 0;1uniformly inx2B; sP0:
Wis uniformly weaklyq-persistent as a consequence of Theorem 3.4. In order to check property
(PS), let us describe the elements ofR t; andR in (2.3) in terms of system (3.4). To this end we
consider sequencessj! 1; yj !B inX, asj! 1: LetIj; Rj be the solutions of
dIj
dt ÿl tsjIa tsj N tsj ÿIjÿRjIjÿc tsjIj;
dRj
dt ÿl tsjRjc tsjIjÿn tsjRj; Ij 0;Rj 0 yj:
After choosing a sub-sequence we may assume that yj converges to some element x. Since N is
uniformly continuous and bounded on 0;1; the Arzela±Ascoli theorem implies that, after
choosing a sub-sequence,
N sjt !N~ t; j! 1
locally uniformly intP0;whereN~ is a bounded continuous function. Moreover, sinceL10;1is
separable, the Alaoglu±Bourbaki theorem implies that, after choosing a sub-sequence,
a sjt !a~ t; j! 1;
c sjt !~c t; j! 1;
l sjt !l~ t; j! 1;
wherea~;c~;l~are elements ofL10;1and the convergence holds in the weak topology carried by
L10;1 as dual space ofL10;1.
The derivatives of Rj and Ij are bounded, uniformly in j2N. Again by the Arzela±Ascoli
theorem we have that, after choosing a sub-sequence,
locally uniformly in tP0; where I~ is absolutely continuous, R~ is continuous, and both are
The same consideration as in the proof of Theorem 3.4 now implies that such an~I cannot exist,
if >0 is chosen small enough, provided thatR~}>1;whereR~}is the analog ofR} in Theorem
Returning to Theorem 3.4 we notice that the weak disease persistence does not depend on the
removal raten. Similarly, checking the proof of Theorem 3.5, we notice that ndid not enter the
proof. This means that the property (PS) is satis®ed uniformly for any continuous non-negativen.
Theorem 3.6. Let N be uniformly continuous and bounded on0;1andR >1:LetNbe an
equi-bounded family of non-negative continuous functions n on 0;1: Then the disease persistence is
uniform inn, in the sense that there exists some >0that is independent ofn2Nsuch thatI1>
for all solutions of (3.1)with I r>0 for somerP0.
Proof.Theorem 3.6 is derived from Theorem 2.5 similarly as Theorem 3.5 from Theorem 2.3. In
order to check condition PSN, we consider sequencessj ! 1; nj 2N; xj !B, whereBis chosen
as in proof of Theorem 3.5. After choosing a sub-sequence we can assume that xj!x2B and
nj !n2N as j! 1; with the latter holding in the weak topology of L10;1. Let Ij; Rj be
solutions of
dIj
dt ÿl tsjIa tsj N sjt ÿIjÿRjIjÿc tsjIj;
dRj
dt ÿl tsjRjc tsjIjÿnj tsjRj; Ij 0;Rj 0 xj:
Arguing as in the proof of Theorem 3.5, elements inR or R t; are given by ~I;
~
I0 tP ÿl~ tI~ t a~ t N~ÿ~IÿR~ ÿ~c t~I t; for almost alltP0;
withR~satisfying (3.6) and~a;~c;l~being the weaklimits ofa s
j;c sj;l sjandN~ being
the locally uniform limit of N sj; after choosing appropriate sub-sequences.
It follows that (PS)N holds, in the same way as in the proof of Theorem 3.5.
Checking the proof of Theorem 3.2 one notices that, under the assumptions of Theorem 3.6, the
convergence in Theorem 3.2 holds uniformly in n2N.
4. The time-heterogeneous S±I±R±S model with distributed removed class
In this section, we model the spread of infection in the same way as in Section 3
N SIR;
dI
dt ÿl tI a tSIÿc tI:
4:1
Again we assume that the disease causes no fatalities and that the population sizeN t is a given
function of timet.l tis the instantaneous per capita mortality rate.c tis the instantaneous per
capita rate of leaving the infective stage.
R t Z a0
0
u t;ada; 4:2
whereu t;is the class age density of removed individuals. Class age is the time that has elapsed
since entering the class.a02 0;1is the maximum possible class age, i.e., the maximum duration
of the removed stage.
One way to model the dynamics of the removed individuals consists in using a partial dier-ential equation
capita rate of leaving the removed stage at class ageaand timet. Autonomous or asymptotically
autonomous versions of this model have been considered by Hethcote et al. [20], Stech and Williams [38] and Castillo-Chavez and Thieme [4].
We make the same assumptions concerning N; l; a and c as in the previous sections. g is
assumed to be a non-negative, bounded continuous function on0;1 0;a0:
The results of the previous section still hold.
Theorem 4.1.Theorems 3.1±3.5hold verbatim.
We brie¯y indicate how the proofs need to be modi®ed and supplemented.
Integrating along characteristics one can transform the boundary value problem (4.3) into an integral equation
This formula can be used to establish existence and uniqueness of non-negative solutions of (4.1)±
(4.3): substituting it into (4.2) reduces (4.1) to an integro-dierential equation inI which can be
solved by Banach's ®xed point theorem. In general, the partial dierential equation in (4.3) will not be satis®ed in a classical sense, but in a generalized sense, which is strong enough to imply
uniqueness. Anyway one can show thatR is absolutely continuous and
we de®ne
n t 1
R t Z a0
0
g a;tu t;ada if R t>0;
n t 0 if R t 0:
Then
dR
dt c tI t ÿl tR t ÿn tR t for almost all t>0
and
n t6 supg 8tP0:
The assertion now follows from Theorem 3.6 and the remark at the end of Section 3.
5. A scalar functional dierential equation
We consider a scalar retarded functional dierential equation that models the growth of a one species population. We assume that state and size of the population couple back to its growth rate
with some delay which does not exceed a number s>0. Let
CCÿs;0; CCÿs;0
denote the space of real-valued continuous functions on the intervalÿs;0and the cone of
non-negative functions, respectively. C is endowed with the supremum-norm and C becomes a
complete metric space under the induced metric.
ForrP0 and/2Cÿs;0 consider
_
x t x tf t;xt; t>r;
x rs / s; s2 ÿs;0; 5:1
where
f :0;1 C!R
is continuous. Here xt 2Cÿs;0 is given by
xt s x ts; s2 ÿs;0:
An example is Hutchinson's [22] equation
x0 t x ta t ÿm tx tÿs; 5:2
wherea is a continuous function andm a continuous non-negative function onR. This equation
models intra-speci®c competition increasing the per capita mortality rate. If competition is not direct but mediated by exhaustion of a vital resource, the density-dependent feed-back is typically
delayed.ais the dierence of the per capita birth and death rates under optimal conditions, while
m gives the increase of the per capita death rate per unit population density. The autonomous
(see [24, Chapters 4 and 9], and the references therein). In this paper, we try to let the time
de-pendence of a and m as general as possible formulating our assumptions in terms of their time
averages.
ar t
1
t
Z t 0
a srds:
Permanence of the population follows if the time averages ofaand mare bounded and bounded
away from zero for larget and r and ifaand m are uniformly locally integrable.
A locally integrable functiona:0;1 !Ris called locally uniformly integrableif
Z t
r
a sds!0; tÿr!0; tPr:
The following result for Hutchinson's equation, (5.2), gives a ¯avor of the general results for Eq. (5.1) (Theorems 5.5 and 5.6).
Theorem 5.1. Letaandmbe uniformly locally integrable and let there existsc0> 0 >0andt0;r0 >
0such that the time averages of aand msatisfy
06ar t;mr t6c0 8tPt0; rPr0:
Then the population is permanent in the sense that there exists somej>1(independent on the initial
data) such that
1
j<lim inft!1 x t6lim supt!1 x t<j
for all non-negative solutions x of (5.2)which are not identically equal to 0.
In particular we have permanence if bothaandmare bounded and bounded away from 0. But
in populations with seasonal reproduction,arepeatedly becomes negative and this is why we use
time averages to come up with realistic conditions for permanence. The proof of Theorem 5.1 (end of this section) checks the assumptions of the results we will now derive for the general scalar equation (5.1).
Assumption 5.2.
(a) We assume thatfis continuous and satis®es the following Lipschitz condition: There exists a
Borel measurable functiong :0:12 ! 0;1such that
jf t;/ ÿf t;wj6g t;k/k kwkk/ÿwk 8tP0; /;w2C:
g t;is non-decreasing for everytP0 andg ;c is uniformly locally integrable for everyc>0.
(b) We assume thatf t;0 is uniformly locally integrable.
(c) Finally, we assume that there exists a uniformly locally integrable functionf~such that
f t;/6f~ t 8tP0; /2C:
In order to show global existence and uniqueness of solutions for initial data inC we extendf
f t;/ f t;/
with / denoting the positive part of /;/ s maxf0;/ sg. Standard FDE theory (for
ex-ample [15,24]) implies that we have unique local solutions of the modi®ed equation. Obviously,
from the form of (5.1), we have that x tP0 for allt>r or x t>0 for all t>r whenever the
respective property holds fortr. Uniqueness implies that
W t;r;/ xt
with x solving (5.1), de®nes a non-autonomous semi¯ow anchored at r0. Assumption 5.2(b)
and non-negativity of solutions now imply that a solution is bounded on every ®nite interval of existence. It follows from Assumption 5.2(a) that the derivative of a solution is integrable on every ®nite interval of existence. So a solution de®ned on a ®nite interval can be extended to the closure of the interval and further extended. This means that solutions exist for all forward times.
We ®rst want to use Theorem 2.4 to derive conditions for the semi¯owWto be point-dissipative
(cf. [16]), i.e., that there exists some c>0 such that
lim sup
t!1 k
W t;r;/k6c 8/2C; rP0:
Proposition 5.3. Let Assumption 5.2 hold. Further assume that there exists a locally integrable
function f1 such that
lim sup
inf/!1
f t;/ f1 t; uniformly in tP0 5:3
and
lim sup
t!1
1
t
Z t 0
f1 sds<0:
Then there exists somej>0 such that
lim inf
t!1 x t6j
for all solution x of (5.1).
By (5.3) we mean that, for everyd>0 there exists some c>0 such that
f t;/6f1 t d 8tP0;
whenever/2C; / sPc 8s2 ÿs;0:
Proof.Choose >0 and t0>0 such that
1
t
Z t 0
f1 sds6 ÿ; tPt0: 5:4
By (5.3), there exists somej>0 such that
f s;/6f1 s =4 8sP0; whenever inf /Pj:
Suppose that the statement of the theorem does not hold. Then there exists a solution xof
_
for some rPs such that
lim inf
t!1 x t>j:
Choosingr large enough we can assume that
x tPj 8tPrÿs:
Hence, for large enought,
x t6x reÿ tÿr=2:
This implies that x t !0 as t! 1 contradicting
lim inf
t!1 x tPj>0:
In order to apply Theorem 2.4 we choose as metric space X C endowed with the metric
induced by the supremum norm. We set
q / / ÿs:
This choice of q will work because, in Assumption 5.2, we have required a global estimate of f
from above, while, via the Lipschitz condition, only a local estimate offfrom below. If it were the
other way round, q / / 0 would be a more appropriate choice.
We turn to the study of the function r in Section 2. In this context, r t;/;s x tsÿs,
wherexis the solution of
_
x t x tf t;xt; tPs; xs /:
Equivalentlyr t;/;s y tÿs, whereyis the solution of
_
y t y tf ts;yt; tP0; y0 /: 5:5
Lemma 5.4.Hypothesis (CA) is satisfied under Assumption 5.2.
_
c0 <1 by Assumption 5.2(c). By (5.1) and Assumption 5.2(a),
_ x t
x t Pf t;0 ÿg t;q xtc0q xtc0:
This means that the following set Bis an absorbing set for the semi¯ow W:
B f/2C; / s6/ rc s;r; / sP/ r~c s;r;q /; ÿs6r; s60g
solution yof (5.5) that
y t6/ 0exp
The ®rst estimate together with Assumption 5.2(c) shows that y t6n t with a non-decreasing
It follows from Assumption 5.2 that the continuity of y does not depend on r and /, so the
Due to the Lipschitz condition in Assumption 5.2(a),
lny t
Theorem 5.5. Let f satisfy Assumption5.2.Assume that there exists a locally integrable functionf1
on0;1with
lim sup
inf/!1
and
lim sup
t!1
lim sup
r!1
1
t
Z t
0
f1 srds<0: 5:7
Then there exist some j>0such that
lim sup
t!1
x t6j
for all solutions x of
_
x t x tf t;xt; tPr; xr /2C;
where rP0; /2C are arbitrary initial data.
Proof. Choosing >0 small enough, it follows from the considerations preceding this theorem
thatyis exponentially decreasing. So the setsR~ jin Theorem 2.4 are empty for all/2X ifjis
chosen large enough. All other assumptions of Theorem 2.4 have been already checked. By
Proposition 5.3, the semi¯ow generated by (5.1) is uniformly weaklyq-persistent and so uniformly
strongly q-persistent. The statement now follows fromq xt x tÿr.
Using Theorem 2.3 rather than Theorem 2.4 we can obtain that the solutions are bounded away from 0. The assumptions of Theorem 2.3 are veri®ed in complete analogy.
Theorem 5.6. Let f satisfy Assumption 5.2andf0 a locally integrable function such that
lim inf
/!0 f t;/
Pf0 t; uniformly in tP0;
lim inf
t!1 lim infr!1
1
t
Z t 0
f0 srds>0:
Then there exist some >0 such that
lim inf
t!1 x tP
for all solutions of x
dx
dt x tf t;xt; tPr; xr/2C;
where rP0; /2C are arbitrary initial data with / 0>0.
Theorem 5.1 follows from Theorems 5.5 and 5.6 withf~ t a t; g t;x m tx, and takingf1
and f0 in the forma t ÿgm t with suciently large or small g>0.
Acknowledgements
Appendix A. Equi-mean-convergent functions
A Borel measurable functionf :0;1 !Ris calledequi-mean-convergentor to haveuniformly
convergent means if the (asymptotic) mean value
Mf lim
t!1
1
t
Z tr
r
f sds
exists, is independent of r and the convergence is uniform in rP0. Obviously the
equi-mean-convergent functions form a vector space. Important examples of equi-equi-mean-convergent functions are asymptotically almost periodic functions and weakly asymptotically almost periodic functions.
A bounded function f :0;1 !R is called asymptotically almost periodic if f is uniformly
continuous and if every sequence tj! 1 j! 1 has a sub-sequence tjk such that f tjk
converges uniformly on 0;1ask! 1.
A continuous bounded function f :0;1 !Ris calledweakly asymptotically almost periodic
if the set of translatesfft;tP0g; ft s f ts; is relatively compact in the weak topology of
the space of bounded continuous functions on0;1(with sup-norm).
The following characterization holds (see [27], or [28], for references, also concerning
gener-alizations to Banach space-valued functions). A bounded continuous function f :0;1 !Ris
weakly asymptotically almost periodic if, for any two sequences sn and tn in0;1,
lim
n!1 mlim!1f smtn mlim!1 nlim!1f smtn;
provided that both iterated limits exists.
Obviously equi-mean-convergence makes sense for locally integrable function while weak as-ymptotic almost periodicity requires the function to be bounded and continuous. In the following we extend the class of equi-mean-convergent functions beyond the class of weakly asymptotic almost periodic functions.
Lemma A.1.A locally integrable functionf :0;1 !Ris equi-mean-convergent if there exists a weakly almost periodic function g such that f ± g is equi-mean-convergent.
The following results are easily checked.
Lemma A.2.A locally functionh:0;1 !Ris equi-mean-convergent to 0 if one of the following holds:
(i) h2Lp0;1for somep2 1;1.
(ii)For every >0 there exist somec>0 such that
Z t
r
h sds
6c tÿr 8tPrP0:
Proposition A.3.A locally integrable functionf :0;1 !Ris equi-mean-convergent if there exists a weakly almost periodic function g such that one of the following holds:
(ii)For every >0 there exist some c> 0 such that
Z t
r
f s
ÿg sds
6c tÿr 8tPrP0:
Obviously the asymptotically almost periodic functions (with values in R) form an algebra
under point-wise multiplication. We conclude with the observation that asymptotic almost
peri-odicity is preserved under convolution with an L1-function.
Lemma A.4. Let f :0;1 !R be asymptotically almost periodic and k2L10;1. Then the
convolution kf,
kf t Z t
0
k sf tÿsds; tP0
is also asymptotically almost periodic.
Proof.This follows from a classical characterization (see Refs. [27,28]). A function is
asymptot-ically almost periodic if and only if there exists a uniquely determined almost periodic functiong
such that f t ÿg t !0 as t! 1.
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