Global Stability and Persistence of Simple Food Chains

H. I. FREEDMAN* AND J. W.-H. Scrtv

Department of Mathematics, University of Alberta, Edmonton, Canada T6G 2GI

Received 27 February 1985; revised 30 May 1985

ABSTRACT

The main purpose of this paper is to develop criteria for which a simple food-chain
model of intermediate type and of arbitrary length has a globally stable positive equilibrium
and to develop criteria under which such a food chain exhibits uniform persistence. The
same techniques are used to obtain conditions for a model of a predator-prey system with
mutual interference of the predator to possess a globally stable positive equilibrium.

1. INTRODUCTION
In this paper we are interested in models of simple food chains of
“higher” order, that is, of arbitrary length. By a simple food chain, we mean
one where there is only one population at each trophic level and where each
population except the lowest eats only the one on the immediate lower
trophic level. Furthermore, we assume that the ecosystem is in a closed
environment with no immigration and emigration of populations allowed.
A large number of food-chain models that have appeared in the literature
are of Lotka-Volterra type. Gard and Hallam [19] consider criteria for
persistence and extinction; Takeuchi, Ada&i, and Tokumaru [34] develop
conditions for the stability of a nonnegative equilibrium; and Harrison [22]
and So [31] derive criteria for global stability of such an equilibrium. In
general, De Angelis and Goldstein [8] give criteria for there to exist a unique
positive equilibrium for Kolmogorov-type food webs.
Most discussions of more general models of food chains have been
restricted to chains of length three. Rescigno and Jones [28] set down
assumptions for a three-dimensional Kolmogorov system to be a food chain.

*Research partially supported by the Natural Sciences and Engineering Research

Council of Canada.
‘Present address, Department of Mathematics and Computer Science, Emory Univer-
sity, Atlanta, Georgia 30322.

MATHEMATICAL BIOSCIENCES 76:69-86 (1985) 69

OElsevier Science Publishing Co., Inc., 1985
52 Vanderbilt Ave., New York, NY 10017 0025-5564/85/$03.30
70 H. I. FREEDMAN AND J. W.-H. SO

Other papers dealing with various aspects of three-dimensional food chains

include: Beddington and Hammond [l], Freedman [ll], Freedman and
Waltman [12,13], Gard [15,16], Saunders and Bazin [30].
For more general food chains and food webs, the reader is referred to
Conrad [6] and De Angelis [7] with regard to questions of stability and
connectance.
In this paper, we will be concerned with the questions of global stability of
an equilibrium and persistence of the top predator population. Using results
in a paper of Butler, Freedman, and Waltman [2], we will show that these
two notions are connected in the sense that the criteria for persistence will
automatically imply the existence of a positive equilibrium.
Global stability is generally not very easy to show, but is considered to be
an important property of an ecosystem (Case and Casten [3]). There have
been results for n-dimensional systems (Harrison [21], So [31], Takeuchi and
Ada&i [33]), but these are of Lotka-Volterra type. For more general types of
systems, there are results only for chains of length two, i.e. for predator-prey
systems (see Cheng, Hsu and Lin [4], Goh [20], Hsu [26]).
In this paper, we extend a technique of Hsu [26] to simple food chains in
order to obtain criteria for global stability of more general food chains. We
note that even in the case of food chains of length three our results are new.
Persistence has been defined by various authors in a variety of different
contexts. An early notion of persistence with regard to ecological systems
appeared in Freedman and Waltman [12]. We refer to that form of per-
sistence as weak persistence and define it as follows: A vector of the form
[x1(%..., x,(t)lT is said to be weakly persistent if for each component
x,(t), kmsup,,., x,(t) > 0. This notion was utilized by Gard [15-181 and
Gard and Hallam [19] in models of food chains. Subsequently, criteria for a
stronger form of persistence was developed in Freedman and Waltman [13].
We refer to this notion as strong persistence and define it as follows:
[x1(t),. . . , x,(t)]’ is said to be strongly persistent if for each component
xi(t), liminf,,+, x,(t) > 0. This notion has been applied to three-dimen-
sional food chains in the abovementioned paper, as well as to competitive
systems (Freedman and Waltman [14]) and to predator-prey systems with
several prey genotypes (So and Freedman [32]). Simultaneous with these
developments, Hofbauer [25] defined what is referred to as permanent
coexistence for hypercycles. This is utilized by Hutson and Vickers [27] to
develop criteria for persistence in a one-predator, two-prey system. The
notions in the above two papers amount to what can be termed uniform
persistence, which we define as follows: A system is said to be uniformly
persistent if there exists S > 0 such that for each component x,(t),
liminf,,,, x,(t)>8>0 for all X=[x,(t),...,~,(t)]~~IntR:. More re-
cently, Butler, Freedman, and Waltman [2] have given conditions under
which weak persistence implies uniform persistence. Here we will use these
GLOBAL STABILITY OF FOOD CHAINS 71

various criteria to obtain conditions which guarantee uniform persistence of

food chains of length n.
The organization of this paper will be as follows. In Section 2, we will
present our model and state some preliminary results. Section 3 will deal with
the global stability aspects of this problem, and Section 4 will deal with
persistence results. We give a four-dimensional example in Section 5.
In Section 6, we will note that Hsu’s theorem can be extended to a
predator-prey model with mutual interference among the predator popu-
lation, utilizing our technique for global stability. Mutual interference was
introduced by Hassell [23], Hassell and Varley [24], and Rogers and Hassell
[29], and the predator-prey model with mutual interference we discuss here
was introduced by Freedman [lo].
A discussion follows in Section 7.

2. THE MODEL
We propose as a model of a simple food chain the system of autonomous
ordinary differential equations

(2.1)
with
d
x,(o)=xi,20, i=l,..., n, . =-$

g(xi) is the specific growth rate of the bottom prey and is assumed to
have the following properties: g: [O,cc) + R; g E C’[O, cc); g(0) > 0, g’(x)
< 0; there exists K > 0 such that g(K) = 0.
Pi(Xi), i=l,..., n - 1 is the predator functional response of the i + 1 th
population in the chain. We assume p, :[0,OD)+ [O,co);p, E C'[O,co);p,(O)
= 0, &(x,)> 0.
si>O and ciao, i=2,..., n, are the specific death rates and food
conversion rates of the ith population.

The above assumptions are consistent with those found in Freedman [ll]
and are sufficient to guarantee existence, uniqueness, and continuability for
all positive time of solutions of the initial-value problem.
At this stage we show that all solutions initiating in the nonnegative cone
are bounded and eventually enter a certain attracting set described below.
72 H. I. FREEDMAN AND J. W.-H. SO

THEOREM 2.1

The set

x2 M
(Xl,...,X,) ER: :Odx,gK,O~x,$--KK---,...,
c2 SZ

M M
04x,+?+ *.. +Xn <K+-+...+- (2.2)
c2 c2 . . . c2 S? S”

where U?: is the nonnegative cone in W” and M = max( x,g(x,) : x1 E [0, K]},
satisfies

(i) .& is positively invariant, and

(ii) forall (x~~,...,x,~)ER:,

(xl(t),...,Xn(t))-+& as t++m

Proof. (i): Let (xIO,..., x,~) E Jai. We will show that

(xl(t),...,Xn(t))EJd forall t20.

Step 1: x1(t) < K for all t > 0. This follows from a standard compari-
son theorem (see Hale [21, Chapter 11) by comparing the differential in-
equality k, Q x1 g( x1) with the differential equation jE1= x1 g( x1).
Step 2: xl(t)+[xZ(t)]/c2 < K + M/s2 for all t > 0. For

M
<Kt- forall t>O.
s2

Similarly, we can prove that for i = 2,. ,n

x*(t) + . . . +- x,(t) M
gK+--•+...+-
M
forall t>O.
x1(t) + -
c2 c* . . . ci s2 s,

(ii): Let (xIO,..., x,~) E R:. We will show that

(xl(t) ,..., x,(t))+& as t++m.

GLOBAL STABILITY OF FOOD CHAINS 73

Step 1: limsup,, +m x,(t) d K. Since the solution of the initial-value

problem

%=x,g(x,), Xl(O) =x10 >, 0

satisfies lim sup, _ + mx1(t) < K, the result follows from a standard compari-
son theorem [cf. step 1 of (i)].
Step 2: limsup,,., xl(t)+[x2(t)]/c2 Q K + M/s2. Let c > 0 be
given. Then there exists a 7’r > 0 such that x1(t) < K + c/2 for all t > Tl. As
in step 2 of (i),

forall t>T,.

Let T, > T, be such that

e-Q’i( K+~+~)-e’z~( x,(T,)+~)~<f forall t>T,.

Then
x2(t) M
x1(t)+-- gK+-•+C forall t>T,.
c2 s2

Hence
x2(t) M
limsupx,(t)+- <K+-.
I’+00 c2 s2

Similarly, we can show that for i = 2,. . . , n

x2(t) Xi(t) M M
limsupx,( t) + - + . ..+ p<K+-+...+-. n
r++CC c2 c2 . . . c, s2 Si

We now consider the existence of equilibria for the system (2.1). Clearly
Eo(O,O,..., 0) and E,(K,O,..., 0) are equilibria. Further, from the properties
of g(xt) and pl(x,) (see Freedman [ll, Chapter 4]), if there exists 0 < xi’)
< K such that pl( xi’)) = s2/c2, and if xp = xj’)g( xj’))/p,( xj2)), then
E,(x;~‘,x$~),O , . . . ,O) is an equilibrium. All other equilibria, if they exist, must
be of the form Ek(xlk) ,..., xlk’,O ,..., 0), 16 k < n, or E,,(x{“) ,..., XL”)),
where all xii) > 0.
74 H. I. FREEDMAN AND J. W.-H. SO

3. CRITERIA FOR GLOBAL STABILITY

In this section we assume that E, as defined in the previous section exists.
For convenience of notation, we relabel En as E*( xj+, . . , x,*). It is the
purpose of this section to derive criteria for E* to be globally stable, i.e. for
E* to be asymptotically stable with domain of attraction the positive cone.
Our technique will be to construct a Liapunov function (see Hale [21,
Chapter lo]) whose domain of validity is the positive cone.
First,fori=l,..., n,ifn-1,wedefine

y(xj)=xi-x,?-x:ln(x,/x:). (3.1)
We note that v( xi) > 0 for xi > 0, x, + x: and v(x:) = 0. Further,

lim K(x~)=~ tm, ~(x,)=+ca.

X,-+ o+

Now K’(xi) =l -(x:/x,) satisfies y’(x:) = 0 and (x, - x:)y’(x,) > 0 for
Xi>O, XifXT.
For i = n - 1, we define

(3.2)

in which case
-S” C,Pn-1(x,-1)
+
v,I-1(%-J=
Pn-1(%-l)
From the system (2.1), we note that - s, + c, pn_ i (x,*_ i) = 0. Then from the
assumed properties of p,_ 1(x,_ 1), it follows that V,_ 1 and V,‘_1 have
similar properties as the other V;‘s listed above.
Finally, we define V( xi,. . . , x,) =C:_lt$(xi), and note that V is a posi-
tive definite function in the positive cone with respect to E*.
We now compute p( xi,. . , x,), the trajectory derivative of V along
solutions of system (2.1):

ti(x I)...) xn)= i I/;‘(Xi)i,. (3.3)

i=l
By substitution and some algebraic manipulations we get
GLOBAL STABILITY OF FOOD CHAINS 75

vy(xi)ii=
i
1-z
iI {xi[-si+cipi_l(xi_l)]-xi+lpi(x;)}

PAX;)
= (xI,- x:.)(-s,+cipi-,(x:_,)-x~+,~ 1 i
+ci(xj-x:)[Pi-l(xi-l)-Pi-l(xE1)]

-~(xi-x:)(xi+~-x~.), i=2 ,...,n-2,

- 3” + cnPn-I(x”-A
y-,( x,_l)jc,_l =
P”-I(x”-I)

x{x”-*[-sn-l + cn-IPn-*(“Al -w-lb,-1))

= [ -48 + %Pn-dxn-1)1

X
Ip”_:;xL_l) n
[-~.-~+c.-~P~-~(x~-~)l-x.]

+ ,‘:;;“x:r:,
[- %I+ c”P”-I(xn-I)l

ok”=
i
1-z
”
I x,[-s”+cnp”_l(x,_l)]

Further,

K--1(x,-l)%-1+ V,‘(x,)k

= [ -s, + c”P”-I<~n-dl

X p _:;*I_,, [-h-1 + c”-IP”-z(L)l -e]

[ ”
%1X,-1
+ S” + Cd-,b-,)I [ Pn-z(~“-2)-P”-*(~n*-*)1~
,.cx n-1) [-

With regard to the above expressions, we note that pi ( xi )/xi, i = 1,. . . , n - 1,

76 H. I. FREEDMAN AND J. W.-H. SO

and x,_ l/p,_ I( x,_ t) are bounded below away from zero and bounded
above for x E A, where JZ’ was defined in (2.2). We also note that g(xl)-
x~(pl(xl)/xl) has a root at x,=x:; each of -s, + c,~,_,(x,*_~)-
x,*+r(p,(x,)/x,), i = 2,. . . , n - 2, has a root at x, = x:;
[x,_~/P~_~(x~_~)][-s~-~ + ~,~tp,~~(x,*_~)]-x,* has a root at x,-t =
x:-t; each of pi(x,)-p,(x:), i=l,..., n -2, has a root at x, =x,T; and
- S, + c,~,-~(x,-~) has a root at xnpl =x,*~~.
From the above comments, we define the functions +, , I);, i = 1,. . , n - 1,
as follows
g(xl)_ “:A(4
=-h-x:)~1(-4,
Xl

pyt’=-(x,-xX:)$,(x,),
- s,+ CiPI4(XL)- x,*+1-
I

i=2 ,...,n -2,

p
n
_:;;‘_
n 1
) [ - s,-1+ -c = -b-1 - X,*-l>
Cn-lPn-2(X,*-2)l
x+n-l(xn-l),
P,W-d-c) = br - a+A
i=l ,.,.,n -2,
-3, + c?lP,pI(x,~A = (T-1 - x,*-J+~~,(~,~,).
(3.4)
We further introduce the notation

u,=x,-xx* I 3 i=l ,...,n. (3.5)

Using the above notation, we can write
n-1
k( Xl ,...,x,) = - c u
IJ
wu
’ J’ (3.6)

where ai, = a,, , a,, =Oif ]i-j]>l,and”‘=’

u,~=$.J~(x~), i=l,..., n-2, an-l.n-1 =~~-*(x,-,)~,-,(x,~-,>~

1
(3.7)
1 Pi(%)
Ur.r+l = - [ ~ - ci+l+i(xt) 9 i=l ,...,n -3 (3.8)
2 5

an-2,n-l = -2
1 Pn-2(&-*) -
[ 4-2
,‘“;;“x”r:,
” n
(Pn-2(Xn-2)+nPn-1(Xn-1)
1
(3.9)
.

Finally, we define A to be the (n - 1) x( n - 1) matrix A = (a,,).

The following is the main result of this section.
GLOBAL STABILITY OF FOOD CHAINS 77

THEOREM 3.1

Let A be positive definite for all X E S? n Int WT. Then E* is a globalb

asymptotically stable equilibrium of the system (2.1) with respect to initial
values in Int R :.

Proof. Let X E & n Int R:. Then, since A is positive definite, 6’(X) < 0.
Theset{X~~~IIntR~:Y(X)=O}isasubsetofS={X~IntR~:xi=
x7, i=l ,...) n - 1). Since the largest invariant set in S is the equilibrium
point E*, therefore, by La&he’s invariance principle (see Hale [21]), E* is
globally asymptotically stable. n

Remark 1. A is positive definite if and only if the principal minors are

all positive. Computing these minors could lead to explicit criteria for global
stability of E* for particular systems (2.1). This will be done in Section 5 for
a three-dimensional system.

Remark 2. A necessary condition for the principal minors of A to be

positive is that aii > 0, i = 1,. . . , n-l.Alsoa,,>Oifandonlyif +i(xi)>O,
0 < xi < K. This can be interpreted geometrically as a requirement that if the
graph of x2 = x,g(x,)/p,(x,) has a local minimum on 0 < xi < K, then xf
must be smaller than this minimum value of the function (see Figure 1). For
a,;>O, i=2 ,..., n-2,then $,(x;)>O, i=2 ,..., n-2. Thisimpliesthat

Pitxi) Pltxf)
if 0 < x, < x,?,
Xi <x:

and the reverse inequality if x: < xi, X E JZ’. Note that this could be
satisfied if pi(xi) is a Holling type-111 (sigmoid) function (see Figure 2).

x2

x9
_-_
--___
-___
k-7
FIG.1. X$ smaller than the local minimum of the function x2 = x,g(x,)/p(x,),
0 < x1 < K, which implies &(xl) > 0.
78 H. I. FREEDMAN AND J. W.-H. SO

FIG. 2. Helling type III (sigmoid) functionaJ response. x, < x:p, (x,)/p, (x: ), x, > XT
only if X 6 a!.

Finally, a,, _ 1,,,_ 1 > 0 implies that J/n-1(x,-1)> 0 [since clearly $J~-~(x,_,)
> 01, and similar statements hold to those in the case i = 2,. . , n - 2.

To conclude this section, we state a corollary which could lead to more

explicit criteria for global stability.
COROLLARY 3.2

Define

a I, 9 i=l,n-1,
ii, =
i 5% 1 i=2 ,...,n -2

ZfforaNXE~nIntR: wehaueii,>O, iiriii+l-af,+l>O, i=l,...,n-2,

then the conclusion of Theorem 3.1 holds.

Proof. The hypotheses of this corollary imply that ii, u,’ + 2a,, , + 1u, u, + 1
+ iii+lu,2-el, i=l,..., n - 2, are positive definite for all X E ~3 n Int BB: . The
corollary now follows from the observation that p can be written as

n-2

ti= - C [ ii,uf +2a,,,+,ujui+l + Zr,+,~f+~].

r=l

Remark 3. We note that in the special case n = 3, the condition on iii in

Corollary 3.2 is equivalent to the condition that A is positive definite in
Theorem 3.1.

4. PERSISTENCE CRITERIA
In the previous section, we have given criteria for there to exist a globally
asymptotically stable positive equilibrium in a simple food chain of length k.
GLOBAL STABILITY OF FOOD CHAINS 79

In the present section we will assume such global stability of equilibria in

order to obtain persistence criteria for simple food chains. We first state three
lemmas which we require in the proof of our theorem.
LEMMA 4.1 (BUTLER AND MCGEHEE)

Let E be a hyperbolic equilibrium in the omega limit set w(X) of an orbit

O(X), where X E R k. Then either w(X) = { E }, or there exist points QS E
WS( E) and Q” E W”(E), where WS( E) and W”(E) are the stable and
unstable manifolds respectively of E, such that QS # E, Q” # E, QS, Q” E
a(X).
Proof See [13, Appendix 11. n

Before stating the next lemma, we introduce the following notation.

Let {iI ,..., ik} C {l,..., n}. Then

*iii,,....;, ={XER::+=O forall i4{i,,...,i,}}.

Note that the Hi,,, ,, i,‘s are invariant sets.

LEMMA 4.2

Let XE *I ,___,i-l,;+1 ,._.,k’ Then o(-VC *I ,..., i-1.

Proof Consider the orbit 0(X) with X as initial condition. Then clearly
[sincep,(O)=O], O(X)cH, ,.,,,, _l,r+l ,,_, ,.Butthe(i+l)thcomponent xi+r
of X satisfies &+t < - siflxi+r, which implies that x,+l(t) + 0 as t + 00.
By induction the lemma follows. n
LEMMA 4.3

Let E, be an equilibrium as defined in Section 2, i.e.,

E, = (xi’),@ ,..., x!“,O ,..., 0). (4.1)
Then the eigenvalues of E, corresponding to eigenvectors in the xi + 1,. . . , xk
directions are respectively -s,+~+c~+~~~(x~~)),-s~+~,...,-s~, i=l,...,k
-1.
Proof. The proof is immediate upon computing the variational matrix of
system (2.1) about E,. n

We now state and prove the main result of this section, for the system

%=xlg(x,)-x*P*(x,)~

Ec/=xj[-si+cjpj_l(xj_,)]-xi+,pj(xj), j=2,...,n-1,

*“=X,[-s”+CnP”-l(Xn-l)l-Xn+lPn(Xn),
%I+1 = x,+1 [ -s,+1+ Cn+lPnGn)l. (4.2)
80 H. I. FREEDMAN AND J. W.-H. SO

THEOREM 4.4

For the system (4.2), assume that E,, i = 1, _.. , n, exist und are hyperbolic.
Further, suppose that E, is globally asymptotically stable with respect to
Int HI .,,.,,. Then if

-sn+1 fCn+lPn (xI;“‘)>O, (4.3)

the system (4.2) exhibits strongpersistence.

Prooj: Let XE Int HI,. ,n+l. We first show that E, 4 w(X), i=

O,l)..., n.
Suppose E, E o(X). Since E, is hyperbolic, { E, } f w(X). Hence by
Lemma 4.1, there exists Qi E I+“( E,)\{ E,} such that Q; E w(X). But
w”(G) = K.....n+i, and all orbits in H2,.,.,n+l \ { E,,} are unbounded,
giving a contradiction, since all orbits in w(x) are bounded.
Suppose now that El E o(X). Since by assumption E, is hyperbolic,
there exists Qf E I+“( El) \ { E,} such that Qf E w(X). By assumption, E, is
globally asymptotically stable in Hl,z, and from Lemma 4.3, - s2 +
c2p1(x{“)> 0; hence W”(E,)= Hl,3 ,,,,, n+l\ H3,,,,, n+l. But by Lemma 4.2,
there exists Ps E HI \ {El } such that Pf E w(X). Since the closures of all
orbits in HI \ { El} either contain E, or are unbounded, we again have a
contradiction.
Similarly, it can be shown that Ez,. , E,_ 1 B w(X). Further, because of
the assumption that E, is globally asymptotically stable in HI, .., , , i = 1,. , n
- 1, no point of HI ,.,,,, can be in w(X). Hence by Lemma 4.2, no point of
H l..... r-1,,+1,...,n can be in 40
Finally, if E,, E w(X), then by (4.3) and the assumption on global
stability, E, is a saddle point. Hence there exists Qi E Int HI,,,,, n \ { E, } =
W( E,)\ { E, } such that Qi E w(X). But cl O( Qi) either is unbounded or
contains points of some HI ,,,,, , _ I, ,+ 1, .., n, once again leading to a contradic-
tion.
Asafinalstepintheproof,wenotethatif o(X)nH, ,,..,, _i,,+i ,,,,, n+lfo,
then there exists P E w(X) such that cl O(P) either is unbounded or contains
oneofthe E,, i=O ,..., n. n

Remarks. Hyperbolicity of E, is equivalent to the requirement that

+ C,+1Pi (x,“)) # 0. (This follows from Lemma 4.3.) The requirement
th:;+z: ,+ 1 be globally asymptotically stable in HI, ..,,+ 1 requires - s, + 1 +
c,+lpi(xl”) > 0.
COROLLARY 4.15

Under the hypotheses of Theorem 4.4, the system (4.2) exhibits uniform
persistence and contains an equilibrium of the type E, +1.
GLOBAL STABILIW OF FOOD CHAINS 81

Proof. The proof is a direct consequence of Theorem 4.4 and the main
theorem of [2]. n

5. A FOUR-DIMENSIONAL EXAMPLE
In order to illustrate the results of the previous two sections, we consider,
as an example of a four-species simple food-chain model, the system

7xY
x=x(4-x)--
4+x’

L=y( -l+g+xp(Yh

i=z[-y+Ep(y)]-wq(z),
ti=ww[-s+cq(z)], (5.1)
where

Y2 + YY Odyd8,
P(Y) = (5.2)
89-17espY, 8<y,

and where q(z) satisfies the general criteria for p, ( xi) given in Section 2.
The four known equilibria for the system (5.1) are E,(O,O, O,O),
E1 (4,0,0,0), E,($, F ,O,O), E3 (3,1,1,0). Clearly El is globally asymptotically
stable in Int Hi, and El is hyperbolic, since - 1 + 7(4)/(4 + 4) > 0.
We define

-1+(7(/4+E)
1/2(x,y) =jx &+Y-:-Tln9&.
213 75‘/4 + 6

Then

t2 = -2(3x+2)(3x-2)2~0
63(x+4)

and E2 is globally asymptotically stable with respect to Int Hi.2. Further,

since - 2 + p (3) > 0, E, is hyperbolic.
For E3, we define

21 Y -2+p(t)
V,(x,y,z)=x-3-31n;+~jl dt+r-1-lnz.
P(4)

After some algebraic manipulations, and utilizing (5.2), i; (x, y, z) can be

82 H. I. FREEDMAN AND .I. W.-H. SO

put into the form

-1
tis(x.y,z) =
10(x+4)(y+1)

x{10(x+3)(y+1)u*- 14(y+7)uu+21(y+4)(y+2)v2),

(5.3)

where

u=x-3, u=y--1.

Since~n{z=0}={(x,y,0):0~x~4,0~x+y~8} for E,,werequire

ti3(x,y,z) c[ 0 on (sfn(z =0})\{(3,1,z)}. Since from (5.3), V,(x,y,z) is
a negative factor multiplying a quadratic form, this will occur provided

196(y+7)2<820(x+3)(y+l)(y+2) (5.4)

on 0 4 x 6 4, 0 < x + y d 8. One can readily verify that the inequality (5.4)

holds in this region, which proves that E3 is globally asymptotically stable in
*nt K,*,3.
Finally, given s and c, if q(z) is such that - s + cq(1) > 0, then the
system (5.1) persists.

6. PREDATOR-PREY SYSTEMS WITH MUTUAL INTERFERENCE

In [lo], a model of a predator-prey system (a food chain of length two)
with mutual interference among the predators was discussed. In this section,
we modify our technique to give a simple criterion for this model to have a
globally asymptotically stable equilibrium for a certain range of one of the
parameters.
The model is given by

(6.1)
where g(x), p(x) have the properties described in Section 2; q(y) is such
that q(0) = 0, q’(y) B 0, and describes intraspecific competition among the
predators for their food; m is the mutual interference parameter, : 6 M < 1.
The lower bound of m is for technical mathematical reasons, and has no
biological significance. m describes intraspecific competition among preda-
tors in searching for prey. Under assumptions given in [lo], we assume a
positive equilibrium E*(x*, y*) exists.
GLOBAL STABILITY OF FOOD CHAINS 83

Since WI< 1, the system (6.1) is not a dynamical system. Using a transfor-
mation described in [9], we set

U=X 9
” = p,
(6.2)
and obtain the transformed system

ir = ug( u) - vm’(i-m)p( u)
Ij = (1- m)[ - sv + cp( u) - “-m”‘-m)q( vi’(i-m))] (6.3)

The system (6.3) is a dynamical system (see [9]) and has a positive equi-
librium ,!?(ii, E), where t = x*, 5 = y*i-“.
We now define F(u) and G(v) by

G(v)=(l-m)ui /(i-m) _ ijm/(i-m)” + m~/(i-m).

(6.4)

It is easy to check that F(u) and G(v) are positive definite functions about
u = t and v = 6 respectively. Hence the function

l’( u,v) = F( u) + G(v) (6.5)

is a positive definite function about B.

We now compute the trajectory derivative ?( u, v) about solutions of the
system (6.3), which after some algebraic manipulation reduces to

qu,v)=(1-m)c[p(u)-p(n)]

m/Cl-m)
_
[
;m/(l-m)
ud u)
po-Cm’“-m’
1
+(1- m)[v 1
x[-su+ cp(ii)-~-m/"-m)q(d"'-m')]. (6.6)

We note that k( u, v) can be written as

ti(u,tJ) =H(u)+L(v).

Clearly H(u) and L(v) have roots of multiplicity two at 5 and E respec-
tively. Hence if H(u) and L(v) are negative definite about “u and 5,
respectively, k and hence E* will be globally asymptotically stable.
H(u) will be negative definite about B provided

( u - ii) [ ug( u) - w -m)p( u)] < 0, uf ii. (6.7)

This is analogous to the hypothesis of Theorem 3.2 in Hsu [26].

84 H. I. FREEDMAN AND J. W.-H. SO

L(u) will be negative definite provided

(u-q[ -su+cp(L)-v--“/‘l~“‘q(u’/“~“‘)] <o, VZiYJ.

(6.8)
This clearly will be guaranteed if su + ~-~/(‘-~)q( ul/(l-m)) is an increasing
function, i.e. if

q’(y)- mY-‘q(Y)‘0. (6.9)

From the above we have proved the following theorem.

THEOREM 6.1

Let i Q m < 1. Let the inequalities (6.7) and (6.8) hold. Then E* is globalb
asymptotically stable for the system (6.1) with respect to solutions with positive
initial conditions.

7. DISCUSSION
It has been shown in previous work (see e.g. Freedman and Waltman [12],
Gard [15]) that food chains can exhibit a variety of behavior, including
persistence or extinction of the top predator, oscillatory behavior, and
globally stable equilibria. In this paper, we give criteria for food chains with
nonlinear functional responses to have a globally stable interior equilibrium,
and criteria for top-predator persistence.
In the case of a globally stable equilibrium, in order for the criteria to be
satisfied, certain of the predator functional responses need to be sigmoid
(Holling type III). Although this is thought to be rare by some ecologists,
there are researchers who believe that this is the natural response for some
populations (see Chua et al. [5] and the references therein).
In the case of a predator-prey system with mutual interference, we again
derive criteria for a globally stable interior equilibrium, thus eliminating all
other possible dynamics (see Freedman [lo]).

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