J. theor. Biol.

(1979) 79, 243-246

LETTERS TO THE EDITOR

A Generalized Prey-Predator Model

The evolution of a community of population consisting of several interacting

species may be described by a set of non-linear equations, a well-known
example being Lotka-Volterra model. The two-species L-V system is
characterized by the property that it is conservative (Goel et al., 1971) and it
possesses centre type of critical point. Though L-V model occupies an
important position in the population dynamics of interacting species, it has
some unrealistic features e.g. the oscillations are initial-value dependent and
this model does not exhibit a limit cycle behaviour.
We have investigated a modification of the L-V model wherein the
interaction term is taken to be proportional to some power of the population
sizes of the two species. This system is non-conservative and does not possess
any centre type of critical point. We have observed, however, that under
certam conditions this model can show a limit cycle behaviour. It is further
seen that when the strength of the self-limiting growth term exceeds a critical
value, the limit cycle behaviour is not observed at all.
For a community comprising of one prey and one predator with
populations N, and N, respectively, we write the evolution equation as

> (1)
dN,
__ = - yN, + /I2 N’fN;
dt
both m and n being positive numbers.
By means of appropriate changes of variables the equations (1) may be
written as
i = ax(l-/!?x)-.xrny” = Xh.i(x,y)
(2)
p = -y+x”y” = YKZ(X, 4’) I
where x and y are the variables corresponding to prey and predator
respectively and the dots represent (new) time-derivatives.
In order that these equations may be biologically plausible, a set of
conditions, known as Kolmogorov conditions (Kolmogorov, 1936; May,
1972) must be satisfied. This requires that
n < 1 and (m+n)> 1.
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0022-5193/79/140243+04 $02,00/O t‘j 1979 .Academic Press Inc. (London) Ltd.
244 GOSWAMI ET AL

The singular point (x,,y,) of the system can be determined from the
following equations

afiXO+xg+n-lM-n) = a
(3)
Yo = axo(l - Pxo) I

The singular point (0,O) is excluded because it is a saddle point.

We introduce the parameters E, 6, n, v and p defined by

& = LY(l-m), 6 = (1 -n), q = a(m+n- I)

v=c&n-2) and p=px,

With the help of usual linearization procedure one can show that the singular
point will be a focus if (E+ v - 6)’ < 4(n - v) and it will be a stable focus or an
unstable focus according as (E + v) < 6 or > 6.
Now, Kolmogorov’s theorem (May, 1972) leads to the result that when
Kolmogorov conditions are satisfied, the singular point is surrounded by a
stable limit cycle provided it is an unstable focus. It is obvious that the region
A in m-n space, which lies inside the parabola (E+ v- 6)’ = 4(n- v) and
above the straight line E+V = 6, corresponds to unstable focus. We must
note that in m-n space Kolmogorov conditions impose two other
restrictions stated earlier, namely n < 1 and (m + n) > 1. So we get another
region B lying below the line n = 1 and above the line m+n = 1, noting that
both m and n are always positive. We have seen that only when the value of p
is less than a critical value there exists an overlap between the regions A and
B. This overlap is the domain of limit cycle.
The main feature of the prey-predator system governed by equations (1) is
the existence of limit cycle for certain values of m and n. It is observed that the
domain of limit cycle is very much altered by the magnitudes of the
parameters a and ~1. Further we see that for a fixed value of cc,the domain of
limit cycle diminishes as p increases.
The existence of limit cycle can be envisaged geometrically as illustrated by
Rescigno & Richardson (1973). It is evident that the phase space is
partitioned into four regions by the curves ~r(x, y) = 0 and x2(x, y) = 0. One
can show, by considering the signs of dy/dx in the four regions, that a phase
space trajectory starting from infinity gradually spirals inward. On the other
hand, for m < 1, n < 1 and (m + n) > 1, the point of intersection of the curves
rcr(x, y) = 0 and K~(x, y) = 0 is an unstable focus. Consequently, a trajectory
in the neighbourhood of the point of intersection will spiral outward (Fig. 1).
Hence the existence of a stable limit cycle is evident.
 LETTERS TO THE EDITOR 245

x /

FIG 1. The behaviour of the phase trajectories, when the critical point is an unstable focus
surrounded by a stable limit cycle (m < 1, n s: 1, (m +n) > 1), is shown.

Finally, we may add another interesting point about this model. When b in
equatxon (2) is zero, then by a transformation
u = log (x” - ly”)
v = log(x”y”-‘)
equation (2) can be written as
ti = a, e”+b, e”+c,
C = a2 e”+b, er+c2 I
These are essentially the Verhulst form of equations occurring in
population dynamics. As shown by Trubatch & Franc0 (1974) one can
construct a linear Lagrangian appropriate to these equations and apply
variational method to obtain the frequency of oscillation.

Depar.tment of Physics DILIP GOSWAMI

Ramsuday College, Amta
Howroh, West Bengal, India

Department of Physics AVUIT LAHIRI

Vidyasagar Evening College,
Calcutta. India
246 GOSWAMI ET AL.

Saha Institute of Nuclear Physics BRAHMANANDA DASGUPI’A

92, Acharya Prafulla Chandra Road
Calcutta-700009, India

(Received 13 June 1978)

REFERENCES

GOEL, N. S., MAITRA, S. C. & MONTROLL, E. W. (1971). Rev. mod. Phy. 43, 231.
KOLMOGOROV, A. N. (1936). Giorn Ins&. Ital. degli Atruari 14, 1.
MAY, R. M. (1972). Science 177,900.
RESCIGNO, A. & RICHARDSON, I. W. (1973). In Foundations qf Mathematical Biolog-v, Vol. III,
Ch. 4, (R. Rosen, ed.). London : Academic Press.
TRUBATCH, S. L. & FRANCO, A. (1974). J. theor. Biol. 48, 299.