PREDATOR - PREY OR PREDATOR POPULATION –

RESOURCE POPULATION

Theoretical Models

The first models to predict the changes in the population

number of a predator population and its resource population

were developed by A.J. Lotka and V. Volterra in the 1920s. In

their equations, they accounted for inverse effect of

population numbers of one species on the other in a predator-

resource system by assuming that the growth rates of a

predator and its resource could be modeled using a

modification of the exponential growth equation.

The basic assumptions of this approach are

1) that the birth rate of the predator is a function of the

number of the resource population and
2) that the death rate of the resource population is a
function of the number of predators.

We will designate

1) the number in the predator population as P and

2) the number in the resource population as R.

First, let us consider the growth rate of the resource

population.
When the predator is not present, then it is assumed that the
resource increases exponentially, or

1
dR = rR
dt

where r is the intrinsic rate of increase for the resource

population.

However, when the predator is present, then the resource

growth rate is reduced by the number of predators and their
ability to obtain the resource.

This reduction can be incorporated into the model as

dR = (r – aP)R
dt

where a is a constant that indicates the successful individual

attack rate by the predators.
If P were constant, then R would grow (or decline)
exponentially.

Because the predator is dependent upon the resource

population as a food resource, when the resource is absent the
predator population declines exponentially.

This rate of decrease can be symbolize as

dP = - dP
dt

where d is the death rate of the predator in the absence of

the resource population.

However, in the presence of the resource, the predator

population will increase at a rate dependent upon
1) the number of the resource population and

2
 2) the ability of the predator to utilize the resource
population.

The growth rate in predator population is then

dP = (- d + bR) P
dt

where b is a constant that indicates the individual conversion

rate by the predator of the resource population into new
predators.
If R were constant, then P would follow the exponential growth

Notice that the last terms in these two equations when they

are multiplied out,

- aPR and bPR, involve the products of the numbers of the two

species. In other words, either the reduction in the growth

rate of the resource or the increase in the growth rate of the

predator due to predation may change very quickly because the

terms resulting in these changes are a function of the joint

densities of the two species.

Let us solve these two equations for their equilibria. The

equation for the growth rate of the resource population can
only be zero if

r – aP = 0

In other words, if

3
P=r
a then dR = 0
dt

and there is no change in the numbers of the resource

population. As we did for two competing species, let us plot this

on a graph that gives the number of predators on one axis and

the number in resource population on the other axis.

In general, the number in the predator population will be larger

than the number in the resource population, so the scales on

these two axes will be different.

The horizontal line in Figure 1 indicates this equilibrium, the

zero isocline for the resource population. Above this line, the

number resource population decreases because of the large

number predators, and below the line it increases because of

the small number of predators.

4
Figure 1: The change in population numbers for the resource species

Likewise, the equilibrium for the predator population occurs

when

-d + bR = 0

because the growth rate pf the predator population is zero.

Therefore, if

R=d
b

then dP = 0
dt

and there is no change in the number of predators. This

equilibrium is called the predator zero isocline and is plotted as
a vertical line in Figure 2.

To the right of this line, the number of predators increases

because of the excess of the resource, and to the left of it

5
the number of predators decreases because there are few
individuals in the resource population.

Figure 2: The change in population numbers for the predator species

At first glance, it would seem that the point where these two
lines cross – that is, where both

dR = 0 and dP = 0
dt dt

would be stable equilibrium. However if the two parts of Figure

1 and 2 are combined, as in Figure 3, then it is obvious that the

numbers of the two species do not approach these equilibrium

densities when they are initially different from these values.

For example, in upper right quadrant, the number of predators

is increasing while the number of prey is decreasing. Likewise,

in the lower left quadrant, the number of predators is

decreasing and the number of prey increasing. As a result, the

6
numbers never converge to the equilibrium but oscillate around

it. The size of the oscilllation is dependent upon the starting

numbers and does not increase or decrease in size with time.

However, random variation in the environment affecting the

species numbers may move them to other predator-resource

combinations. The numbers will then cycle from this point,

indicating that the cycles do not converge to a single stable

cycle. However, the equilibrium numbers do indicate the

average expected number of each species over an extended

period of time.

Figure 3: The joint change in numbers for the resource and predator populations

7
The numbers of the two species can also be plotted over time.

Figure 4 gives the population numbers for both species for two

cycles, beginning at a point in the middle of the lower section

of Figure 3.

Notice that the predator numbers follow the resource

numbers, reaching maximums and minimums a quarter of a cycle

later. The differences in the number of the two species results

from a time lag effect resource numbers on predator numbers.

For example, in lower left quadrant in Figure 3, resource

numbers are increasing and predator numbers are still

declining, and the predator numbers do not begin to increase

until the lower right quadrant.

Let us illustrate these changes with a numerical example.

Assume that r = 0.2 and a = 0.01, so that the number of

predators at which dR/dt = 0, the resource zero isocline, is

P = r/a = 0.2/0.01 = 20.

Likewise, if d = 0.2 and b = 0.001, the resource number at

which dP/dt = 0, the predator isocline, is

R = d/b = 0.2/0.001 = 200.

8
If we let P = 10 and R = 100, a point in the lower left quadrant,
then the change in resource numbers is

dR/dt = (r –aP)R = [0.2 – (0.01)(10)]100 = 10.

The change in predator numbers is

dP/dt = (-d + bR)P = [-0.2 + (0.001)(100)]10 = -1

Therefore, the number in the resource population is increased

to 110 and the number of the predator population decreased to

9. Similar change can be calculated for points in the other

three quadrants. One prediction of this simple model is that

predator and prey numbers should be oscillate.