Nt

0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15

10
19
34.39
56.9533
81.4698
96.5663
99.8821
99.9999
100
100
100
100
100
100
100
100

The model of lo
(2.7):

Rm
K
N0

Number of individuals, Nt

1
100
10

Here, Rm is the m
assumption that
competition for a
the environment
there is no more

120
100

Using this simu

will show:

80
60

asymptot

40

damped s

20

neutrally

0
0

10

15

Time, t

How was this sheet programmed?

The formulas in the columns A and B were constructed in the same way as in
the case of the exponential growth, only the increment boldfaced below is
now different, consistently with the equation 2.7. Thus, in cell B3 we have the
formula: =B2+$E$7*(1-B2/$E$8)*B2.
Note that the references to the cells E7 and E8 contain dollar signs, as these
are model parameters within the yellow-highlighted region and we do not wish
these to be changed when the formula is copied downwards.
Note that for low values of Rm the numbers of individuals, predicted by
this model, are approaching an equilibrium value the carrying capacity, K.
When Rm > 1, but close to 1, you may not notice from the graph that the
approach to the equilibrium is oscillatory. Look, however, at the values in
column B! You will find some, even if small oscillations.
Try to set Rm equal to some large value, e.g., Rm =5. The model will predict
negative numbers of individuals. To get rid of this problem, a modification of
the logistic equation, analyzed in the subsequent sheet Modified logistic
is often used.

displacem
unstable,

The model of logistic growth of the population is defined in the main text in equation
(2.7):
N t N t 1 Rm (1 N t 1 / K ) N t 1 .

(2.7)

Here, Rm is the maximum individual rate of increase (maximum value of R under the
assumption that there is a very low number of individuals in the population, so that
competition for a scarce resource does not yet occur, and K is the carrying capacity of
the environment - the population density where all living space is fully utilized and
there is no more room for additional growth.
Using this simulation model, solve Exercise 2.2. Then confirm that the population
will show:
asymptotic approach to equilibrium when Rm < 1,
damped stable oscillations when 1 < Rm < 2,
neutrally stable oscillations with amplitude determined by the initial
displacement when Rm = 2,
unstable, often non-periodic behavior when Rm > 2.

Back to the introductory text

Nt
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15

10
24.596
52.2806
84.2526
98.6219
99.9904
100
100
100
100
100
100
100
100
100
100

Model:
N t N t 1e
Rm
K
N0

Number of individuals, Nt

In some cas
biologically m
population si
the logistic eq

Rm (1 N t 1 / K )

1
100
10

120

Because the

100

e Rm (1 N t 1 / K ) is

80

We sugge
initial numbe
and see, how

60
40
20
0
0

10

15

Time, t

How was this sheet programmed?

The formulas in the columns A and B were constructed in the same way as in
the previous sheet (Discrete logistic), only the formula is now different,
consistently with the equation 2.7a. Thus, in cell B3 we have the formula:
=B2*EXP($E$7*(1-B2/$E$8)).
Note that the exponential function, ex, is expressed in EXCEL as EXP(X).
Note that for any, even very large values of Rm, the numbers of individuals,
predicted by this model, are always positive. Even if the graph may suggest
these values are equal to zero, the values in column B are always larger than
zero. Effectively, however, the biologist would consider such population as
extinct.

the p
reason
the in

with
equili
oscilla
critica
beyon
non-p

In some cases the logistic growth described by equation (2.7) does not give
biologically meaningful predictions for some large values of Rm is the predicted
population size, Nt, negative. To get rid of this problem, the following modification of
the logistic equation is often used:

N t N t 1e Rm (1 N t 1 / K ) .

Because the regulatory term Rm (1 N t 1 / K ) is now in exponent, the value

e Rm (1 N t 1 / K ) is never negative and so is the resulting prediction on Nt.

We suggest you to run the simulations of equation (2.7a) for various values of the
initial numbers of individuals, N0, and for various values of the parameters Rm and K
and see, how the systems behaves in response to their changes. You will find out that:
the predictions of the number of individuals are always biologically
reasonable, i.e., positive, independently of the choice of the initial numbers of
the individuals and of the parameters;
with increasing Rm, the system predicts first asymptotic approach to
equilibrium, then damped stable oscillations, followed by neutrally stable
oscillations. For which value of Rm does this occur? Try to calculate this
critical value first, before you perform the simulations! When Rm increases
beyond this critical value, we can observe unstable oscillations and finally a
non-periodic solution, which is in biological literature known as chaos.

way as in
different,
formula:

X).
ividuals,
suggest
ger than
ation as

(2.7a)

Back to the introductory text

Nt

0
30
60
90
120
150
180
210
240
270
300
330
360
390
420
450
10
10
50.5309
50.5309
123.104
123.104
81.2197
81.2197
113.887
113.887
88.6986
88.6986
108.708
108.708
92.9366
92.9366
105.537
105.537
95.526
95.526
103.537
103.537
97.1505
97.1505
102.263
102.263
98.1808
98.1808
101.449
101.449
98.8372
98.8372
0
150

0
105.763
123.266
107.75
83.7212
60.9854
42.647
28.9945
19.3103
12.6597
8.19712
5.25454
3.34044
2.10886
1.32347
0.82634

Model:
N t N t 1e Rm (1 N t 1 / K )
Rm
K
N0

Number of individuals at
time t, Nt

N t-1

0
50.5309
50.5309
123.104
123.104
81.2197
81.2197
113.887
113.887
88.6986
88.6986
108.708
108.708
92.9366
92.9366
105.537
105.537
95.526
95.526
103.537
103.537
97.1505
97.1505
102.263
102.263
98.1808
98.1808
101.449
101.449
98.8372
98.8372
100.928

140
120
100
80
60
40 0
20
0
0

100

1.8
100
10

200

300

400

500

Number of individuals at time t-1, Nt-1

Comparison of the discrete logistic model N t N t 1 Rm (1 N t 1 / K ) N t 1 with the m

logistic model,

N t N t 1e Rm (1 N t 1 / K ) .

In this sheet the two logistic models that were analyzed in the previous sheets are compared.
lines in the figures above show, how the number of individuals at time t, Nt, depends on the n
individuals at time t -1, Nt-1. The shapes of these curves are visually similar, but the curve on
(that represented by equation 2.7) predicts negative numbers of individuals at time t, if the n
individuals at time t 1 is large. This does not occur in the figure on the left. The green bro
represent situation, when Nt = Nt-1. Thus the intercepts of the blue and green lines represent
of the system the situation, when the numbers of individuals do not change in time.
The red lines show, how the numbers of individuals change in time. We start on the
the position of the red circle, which denotes the number of individuals at the old time 0. Th
continues vertically to its intercept with the blue line the y-coordinate at this instant
prediction of the population size at the new time 1. Then the red line continues horizontally
intercept with the green line. At this point, the x- and y-coordinates are the same. We can no
the old value by the new value and continue again vertically, to find the predicted popul
at the following instant, time 2. We find it again as the intercept of the red line with the blue l
on.
We do not explain here, how these figures were obtained the explanation would
complicated and is not important.
0
150

Nt
0
0
30
82.8
60
151.2
90
205.2
120
244.8
150
270
180
280.8
210
277.2
240
259.2
270
226.8
300
180
330
118.8
360
43.2
390
-46.8
420
-151.2
450
-270
10
0
10
29.2
29.2
29.2
29.2
80.7789
80.7789
80.7789
80.7789
190.135
190.135
190.135
190.135
281.194
281.194
281.194
281.194
211.021
N t 1 / K ) N211.021
t 1 with the modified
211.021
211.021
276.824
276.824
276.824
276.824
217.42
217.42
217.42
revious sheets are compared. The blue
217.42
274.089
at time t, Nt, depends on the number of
274.089
274.089
ually similar, but the curve on the right
274.089
221.269
individuals at time t, if the number of
221.269
ure on the left. The green broken 221.269
lines
221.269
272.127
ue and green lines
represent equilibria
272.127
o not change in272.127
time.
272.127
223.956
ange in time. We start on the x-axis,
at
223.956
iduals at the old time 0. The red223.956
line
223.956
270.618
y-coordinate at
this instant represents
270.618
270.618
ed line continues
horizontally, until
its
270.618
225.982
ates are the same.
We can now replace
y, to find the predicted
size
225.982 population225.982
of the red line 225.982
with the blue line and
so
269.403
269.403
269.403
ned the explanation
269.403 would be rather
227.585
227.585
227.585
227.585
268.396
0
500

Number of individuals at
time t, Nt

N t-1

Nt1Rm(1/K)
Rm
K
N0

500
400
300
200
100
0
-100 00
-200
-300
-400
-500

100

2
250
10

200

300

400

Number of individuals at time t-1, Nt-1

Back to the introductory text

0
500

500

Nt1Rm(1/K)