Math1 Comput. Modding, Vol.II,pp.25C252, 1988 08957177/88$3.00f0.

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POPULATION MODELS II

SATURATION PROCESSES'

C. 0. A. Sowunmi

University of Ibadan, Ibadan, Oyo State, Nigeria.

Abstract. Saturation, an important occurrence in 'species'

interaction, is given an abstract mathematical formulation which
enables the qualitative study of the dynamics of such interactions,
without approximation, and the consequent a priori restriction of
analysis to some bounded region of the state space. A
qualitative study of Predation illustrates the ideas.

This paper is in two parts:

I. A general formulation of saturation processes;
II. Application to Prey-Predator Interaction.

IKeywords: Predation, Saturation, Species interaction, Stability,

Ultimate boundedness.

INTRODUCTION Then

1.
t t
A GENERAL FORMULATION OF SATURATION B(t;ri) = Fi([slll,.-.,[snln) (1)
PROCESSES
where [.I. is a functional on the history
There are many processes whether natural Of Sj Up I 0 time t. Since no 'birth'
or artificial which essentially involve the is possible in the absence of anyone of
interaction of a number of species, the the species Sl,...,Sn, it follows that
outcome of which is the production of some
species which may or may not be the same Fi(...,O,...) = 0 (2)
as the interacting species. Examples are:
chemical.reactions, bisexual reproduction, If Fi is sufficiently differentiable in the
vector-borne disease propagation etc. In neighbourhood of the origin in its domain,
many of such interactions it is quite usual then
for the production rate to depend directly i
on the concentration/density of each of the B(t,r$ = al. &;I>... Q$+ (3)
interacting species but in such a manner p;.--p: 1
that if the concentration of one of the terms of order higher than pi + . .. +
interacting parts were to be fixed, the p;; ,i. being a positive number, and
production rate would rise with the p+...p,i
concentration/densities of the rest only
up to some maximum value. Furthermore, cy=,p; > 0.
this maximum value increases with the
iCS,,... '5k...6,U) = [Fi(6,0... 6k...6nO)
fixed value of the one component. It is
this rising and levelling off which is here
- Fi(61U... 6kO... 6,+1/(6k-6kO) 2 0 (4)
referred to as Saturation (Sowunmi, 1986).
$(610...6k...6n0 + 0 (5)
Now for a formal definition of the process.
as 6 + m for arbitrarily fixed
Saturation Processes (Deterministic) k
MlD.._Q)..6nU). If
Let S l,...,S, be n different species, with
limFi(d10...6k...6no) = fi(6k) (6)
densities sl,...,sn respectively, functions
of the real variable t20. Let all the
as 6 + m for allr %k , then fi(6k) is
species be involved in an interaction r0
whose resultant is the m species Rl,.._,R,, monotone increasing with 6k. It follows
with densities rl_..,r,, also functions
from [1.4-1.61 that
of time t. Let the expected density of
new members of Ri at time t be B(t;ri).

250
 Proc. 6th Int. Co& on Mathematical Modelling 251

B(t;ri) 2 min[fi(bl),...,fi(6,)1 (7) L2(x,y) = 0 meets only the x-axis at

positive solutions x1 of s-xlT(xl,O) = 0.
where 6i = [s;li. This last inequality can
be regarded as the generalized law of the Gradient of the curve L2(x,y) = 0 is
given by
minimum (Terrien et al., 1957).
r+d
x.
y-xr
Y
II. PREY-PREDATOR INTERACTION WITH Since g has property (4),TtxTxZO, hence a
SATURATION
sufficient condition for x to be unique is
that the gradient of the c&e L =0 is
Let x(t),y(t) be the prey and predator
non-negative. Furthermore a suf P,
icient
populations respectively at time t and
condition for L =0 and L =0 to intersect is
subject to the following pair of equations 1 2
that x <a and the gradient of L2=0 is
f = ax - bx' - f(x,y) 1 b.
(8) non-negative, i.e.
y-xTy(x,y)>O everywhere.
y = EY - yy2 + g(x,y) (9)
The equilibria can be analysed in the
where a,s'O;b,y>O;x,yZO. The variables f,g usual manner, but we shall consider only
are the interaction terms: f(x,y) is the (5,~) locally. Linearising equation (13)
prey death rate due to the predation about (S,rl) yields
only, whilst g(x,y) is the predator renewal
rate due to the predation only. By
application of (2) to f and g, assuming at (15)
least C2 smoothness, we have
Analysis of equation (15) by a well-known
f(x,y) = xyQ(x,y) and
method leads to the following result:
g(x,y) = xYr(x,Y) (10)
Theorem 1
and by (4)
Suppose that the per capita differential
between the predator conversion rate and
@(x,y) + YQy(X'Y)>O
its population stress factor, i.e.
(XT-yy)is always falling relative to the
@(x,y) + x@x(x,y)>O
predator population. If the equilibrium
(S,n)exists, it is locally stable
l-(x,y) + Yry(x'Y)tO
provided that the per capita death rate
of the prey (bx+yQ) is rising relative to
r(x,y) t xrx(x,y)'o. (11)
the prey population; it is otherwise
unstable. Furthermore, if there is
When equality holds in any of (11) we shall
saturation at (S,n)it could be a focus,os a
say there is saturation.
node but not a spiral sink.
1n applying (6) we shall assume that the
Next we shall investigate the ultimate
f;s are linear homogeneous, specifically boundedness of the system (13).

f(x,y)<min[k,y,xl,g(x,y)'min[k2y,k3xl,
I
(12) We first establish the continuability of
every solution of (13) on the whole of
where kl,k2,k3>0 and are constants. Rf. From the first equation in (13),
Thus (8) and (9) become
2 (xe-at ) =-e*t(bx2+f(x,y)). (16)
B = x[a-bx-y@(x,y)J
Therefore
y = y[-c-Yy+xr(x,y) 1 (13)
at
x(t)=xoe -

J’
Equilibrium points are (O,O), (E,O), and
possibly (E,n) the simultaneous solution(s) ea(+'){bx2 (T) + f(x(T),y(T))}dT (17)
of the pair
0
Ll(S,n)~a-bS-nQ(S,n) = 0 where x0=x(O). Similarly,
L2(S,n)EE-YTl-SrK,n) = 0, (14)
y(t) = yoe-ct +
J8te-E(t-T'
if such exist.
x{-yy2 (~1 + g(x(T),y(T))}dT, (18)
The curve L1(x,y) = 0 meets the x-axis at
where y. = y(O). Since the integrand in
(g,O) and the y-axis at positive solutions (17) is non-negative,
y1 of a-yl@(O,yl)=O if they exist; whilst at
x(t)< xoe (19)
252 Proc. 6th Int. Conf. on Mathematical Modelling

Using this last inequality in (18) gives COMMENTS

--Et
y(t) lyoe + jote+-')g(xOe%'(r))dr~20) The concept of Saturation is formalised in
part I of this paper and applied in Part II
to a simple model of prey-predator
The second condition (12) interaction. The interaction terms of the
imposed on g implies model equations replace the classical
mass-action terms. The former are more
g(x,y)<mintk3x,k2yl<fIk3x + k2yl, (21) realistic than the latter yet the resulting
problems are not less tractable. There is
since the terms involved are positive. every likelihood that the concept will yield
Hence, meaningful results in applications to other
t e-E (t-T) areas like disease transmission, bisexual
-Et + 1
y(t)<=yoe population dynamics, etc.
i0 2
(k3xOe aT'+ k2y(T))d-r. (22) Acknowledgement

The author's participation in the Sixth ICMM

It is clear, by the application of
was due to financial assistance notably
Gromwall's inequality to the above, that
from National Oil and Chemical Marketing
y is bounded on any closed finite interval
CO. Ltd., Nigeria and Health Aids Ltd.,
in R+. Boundedness of x follows of course
Lagos, Nigeria.
from (20). Continuability of every
solution on R+ is thus established.

REFERENCES
To investigate the ultimate boundedness of
(x,y), we shall make use of the well-known
La Salle, J. and S. Lefschetz (1961).
result (La Salle and Lefschetz, 1961),
Stability by Lyapunov's Direct Method
theorem XVII. We need a real valued
with applications. Academic Press,
function V(x,y) which is C1 -continuous,
London, P. 118.
such that V(x,y)-tm as \I(x,y)I\-. If we
can show that V(x,y)f-GtO for all (x,y)
Sowunmi, C. 0. A. (1986). A Mathematical
outside some comparct set M, then the
Essay on Polygamy and Mass-Action.
system (13) will be ultimately bounded.
(An Inaugural Lecture delivered at
The choice V(x,y) = x+y;x,ytO meets
the University of Ibadan) (to be
the requirements. Furthermore
published).
Q(x,y) = x+y = ax-bx'-f(x,y)-cy-yy*+g(x,y)
Terrien, J. G. Truffaut, J. Caries (1957).
1 Light, Vegetation and Chlorophyll.
'ax-bx'-my-yy2 + p3x + &,y
Hutchinson Scientific and Technical,
London, pp. 92, 114, 130, 198.
from (21) and the non-negativity of f.
Hence

+(x,y)<=-6< 0 if
k3
(at-)x - bx2C-6 and
2 2
k2 - s)y-yy2< -5.
(_
2 2

Hence M can be taken as the compact set

2a+k3+&(2a+k3)*+8b&
X2

4b
k2-2s+J(k2-2#+8b6
y< 1. (23)
4Y

Theorem 2

Provided that b,y>O the system (13) is

ultimately bounded.