MATHEMATICAL

293

BIOSCIENCES 20, 29%291 (1974)

Exact Solutions of Lotka-Volterra

Equations

MOSTAFA A. ABDELKADER
25 Sh. Chdmpollion, Alexandria, Egypt
Communicated

by Richard Bellman

ABSTRACT
The Lotka-Volterra
nonlinear differential equations for two competing species P
and Q contain six independent parameters. Their general analytic solutions, valid for
arbitrary values of the parameters, are at present unknown. However, when two or
more of these parameters are interrelated, it is possible to obtain the exact solutions in
the P, Q phase plane, and six cases of solvability are given in this paper. The dependence
of the solutions on the parameters and the initial conditions can thus be readily
investigated.

The well-known
competing species
;

Lotka-Volterra

= (1. - UP -

bQ)P,

differential

z=

equations

(s - CP -

for

two

dQ)Q,

with the initial conditions

P(0) = P, and Q(0) = Q,,, and where the
parameters a, b, c, d, r, and s are positive numbers, do not seem to have
been integrated
exactly. In view of the six independent
parameters
in
Eqs. 1 and the imposition of arbitrary initial conditions, the use of computers for their approximate
integration,
for arbitrary
choices of the
values of these parameters and initial conditions,
can give only a limited
picture of the totality of possible solutions.
The exact analytic solutions of these nonlinear equations for arbitrary
values of all the parameters must be quite complicated expressions, as will
be shown below, and they remain at present unknown.
However, when
two or more of the parameters are interrelated,
the writer has been able
to find the exact solutions, and it is the aim of this note to give these
solutions. The solutions involve a variety of algebraic, transcendental,
and
Bessel functions, as well as exponential
integrals, and the dependence of
the solutions on the values of the parameters and the initial points can be
0 American Elsevier Publishing Company, Inc., 1974

294

MOSTAFA A. ABDELKADER

the relative importance

of certain
readily investigated
(for instance,
parameters,
and the changes in the solution curves as these vary, can be
easily assessed).
We shall give here the solutions in the P, Q phase plane, that is, the
one-parameter
families of solutions of the differential equation
dP

(r - UP - bQ)P

(2)

dQ=(s-cl-dQ)Q

[obtained from Eqs. 1 by division] when some of the parameters

are
interrelated.
The time dependence of P and Q, when the solution 4(P, Q)
= C is known, may be obtained if one replaces Q or P in the solution by
1
i
respectively, to get a differential equation for P or Q with the independent
variable missing. In one of the following cases we shall give P(t) and Q(t)
explicitly.
It is obvious from Eq. 2 that the case for which
R = Ir - ksl + Ia - kc1 + lb - kdl = 0,
where k is a nonzero constant, is of no interest, and so we assume throughout that R > 0. We have the following six cases.
CASE

I
s =

The general

solution

r.

of Eq. 2 is given by

b-d+(a-c);

where c = P/Q, C is a constant

= exp

+rju&=c,

(c-

of integration,

2a)[ -

(3)

and
b

(b - d)[ + (a - c)c2 dr*

The solution (3) has different

vanishes or not; thus,

forms

accordingly

as (a - c)(b - d)

(i) for (a - c)(b - d) # 0, Eq. (3) reduces to

a+1
Pp+lQ--p (a - c); + b - d
1
+

rJiq(a - c)l:+ b -

d) d5 = C,

(4)

EXACT

SOLUTIONS

OF LOTKA-VOLTERRA

295

EQUATIONS

where
b

a=

=d-b

a(2d - b) - cd
(a - c)(b - d).

The solution 4 depends to a large extent upon the values of p and a,

which determine the expression for the integral. For instance, if p = - 2
and a = - 3, so that 2d = b and 4c = a, we get the transcendental
solution

z-l

- cr log
Jzz

- J3 cr arctan2j:

= C,

+ z + 1

where
3cP

z=

.
i dQ+l
In contrast, if either p or a is a positive integer, the solution
of algebraic curves.
(ii) For c = a, the solution

is a family

3 becomes

(b - d)PrdQ-7b exp {r&/Q}

+ YJP exp

(74)

d[ = C,

(5)

where T(d - b) = 1.
(iii) For d = b, Eq. 3 becomes
(a - c)PKCQKO exp {IcbQ/P} + r J ivexp

{~b/i}

dc = C,

(6)

where
7c(a - c) = 1,

v(a - c) = c - 2a.

According to the values of the relevant parameters,

the integrals in
Eqs. 5 and 6 can be evaluated in terms of either the exponential function
or the exponential integral.
CASE

II

-r =
The general solution

d(a - c)

a(b)c*o,

a>.

of Eq. 2 is

b-dQ
_ c
PQ = C P + a

where
a = d(c - a),

z = a@ - d),

v = bc - ad.

296

MOSTAFA

A. ABDELKADER

CASE III
d = b.

c = a,

The general

solution

of Eq. 2 is
P +

%Q
+ CP'_"Q" = 1,

where a(r - s) = I.
We get
p(t) =

; +

!$

-1

@-*Jr

C e-

and
Q(t) = $P(t)

ecserjt,

where C is given by
?Po +

;Q,+ CP;-"Q; = 1.

We notice that if s > r, we have P(m) = 0 and Q(co) = s/b; while if

= r/a and Q(a) = 0. All of these ultimate values are
independent
of the initial values.

r > s, we get P(m)

CASE IV
n
s
-==
I
b + d

The general

solution

2ad
C=bdd.

of Eq. 2 is

aP2 + {(b + d)Q -

r}P

= CQ,

where 0 = I + b/d.
CASE V

2s = r,
The general

solution

2c = a,

d = 2b.

of Eq. 2 is
-cl;(z)

aP = bQ + JarP

Cl*(z)

+ Z&(z)
+ Z_+(Z)

where 3bQz = Vat-P , I, denotes the modified Bessel function of the first
kind of order v, and primes denote differentiation
with respect to z.

EXACT
CASE

SOLUTIONS

OF LOTKA-VOLTERRA

EQUATIONS

VI
s =

The general

solution

2r,

c = 4a,

Sd = 26.

of Eq. 2 is

2A + B + 51
2A - B -5

where
A = JSbrQ,

B = bQ - 5aP.

297