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It sometimes seems that there are daily news reports about the threats of global
warming leading to the extinction of species. However, most of the introductory
treatments of models for population dynamics in mathematics classes typically consider
only population growth models. In this article, we will construct a pair of more general
models that encompass the possibility of both growth and extinction.
Let’s think about the behavior of the solutions in each of these regions. In Region
IV, where P < 0, we should expect that the solutions decay toward -∞. In Region I, where
P > L, we should expect that the solutions decay toward L, as we have with the logistic
model. Similarly, in Region II, where K < P < L, we should expect that the solutions rise
toward L, eventually in an asymptotic manner, much as they behave with the logistic
model. Finally, in Region III, where 0 < P < K, we should expect that the solutions
decay toward zero.
But, what should be the actual pattern for the decay in Region III? Should it be a
purely concave up pattern, as with exponential decay? Picture what happens in the
logistic model with solutions that start on either side of the equilibrium level L. If the
initial population is significantly above L¸ the solution will drop very rapidly at first and
eventually slow down as it approaches L asymptotically. If the initial population is very
close to, but above, L, it will decay very slowly toward L. If the initial population is
equal to L, it will remain at that level indefinitely. If the initial population is slightly
below L, and above the inflection point at ½L, it rises slowly in a concave down pattern
as it approaches L asymptotically. Finally, if the initial population is well below L, and
also below the inflection point, it rises ever more rapidly until it passes the inflection
point and then begins to slow. The change from one behavior pattern to the next happens
continuously.
Consequently, we should expect that, in the
extended logistic model we are trying to create, there
should also be a continuous change in the behavior of
solutions depending on where they start with regard to
the new equilibrium level at the minimum sustainable
population. In particular, the assumption that the
behavior of the solutions in Region III automatically
follows an exponential decay pattern seems
inappropriate since that presumes a sudden drop in the
population if the value is slightly below K. Rather, as Figure 4
shown in Figure 4, it makes more sense to expect that the solutions will display a
vertically reversed logistic shape and that there will be an inflection point level in this
region as well.
We show the right-half of the t-P plane in Figure 5 along with the signs of P’ or
∆Pn needed to produce the patterns of behavior we would expect in each region. In
particular, we need to analyze the signs of the factors needed for either P’ or ∆Pn in each
of the four regions that will lead to these behavior patterns. Clearly, in Region I, we need
P’ < 0 or ∆Pn < 0. In Region II, we need P’ > 0 or ∆Pn > 0. In Region III, we need P’ < 0
or ∆Pn < 0. Finally, in Region IV, we need P’ < 0 or ∆Pn < 0.
3( K + L) � 9 K 2 - 14 KL + 9 L2
P= .
8
Unfortunately, this expression provides little P'
P
1.00E+08
insight into the locations of the inflection points. -200 300 800 1300 1800
-4.00E+08
-9.00E+08
-1.40E+09
-1.90E+09 Figure
Figure 6 6
To gain a better feel, let’s consider an example with specific values for the parameters.
Suppose that the population in question has a maximum sustainable level of L = 2000 and
a minimum sustainable level of K = 200. With these values, the above quadratic formula
gives P = 131.73 and P = 1518.27 as the two roots, correct to two decimal places. We see
that the first inflection point corresponds to a height of roughly ⅔ of the minimum
sustainable population and the second inflection point corresponds to a height of roughly
3/4 of the maximum sustainable population. We show the graph of the corresponding
cubic function of P’ versus P in Figure 6 without the –α coefficient.
800
400
0
0 5 10 15 20 25 30
equation for Pn+1 become extremely large very quickly; rather than falling into any
reasonably smooth pattern, the successive values typically jump from one of the four
regions far into a different region and may bounce around dramatically.
Suppose we then select α = 10-10 and again use K = 200 and L = 2000, as above.
We show the results of starting with initial values of P0 = 500 and P0 = 1200 in Figure 7.
Notice that both curves approach the horizontal asymptote of 2000, which is the
maximum sustainable population level, very quickly – the upper curve starting at P0 =
1200 essentially reaches it in about n = 8 time periods and the lower curve starting at P0 =
500 gets there in about n = 28 time periods.
Figure 7
Figure 7: Solutions starting from P0 = 500
and P0 = 1200
-200
Figure 9
roughly P = 132. The solution starting at P0 = 75 begins below this level and its shape
follows a decreasing, concave up pattern. In contract, the solution starting at P0 = 180
begins above this level and its shape is obviously decreasing and concave down at first,
but clearly changes to decreasing and concave up at around n = 250, where P is roughly
130, as predicted. Finally, the solution starting at P0 = -50 is obviously decreasing and
concave down in the view shown; beyond about n = 250 it drops extremely rapidly as it
decreases toward -∞.
References
1. Allee, W. C. (1931). Animal Aggregations. A Study in General Sociology. University of
Chicago Press, Chicago. ISBN 0-404-14501-9.
2. Allee, W. C. (1949). Principles of Animal Ecology. W.B. Saunders Co., Philadelphia.
ISBN 0-7216-1120-6.
3. Clark, Colin W. (2005). Mathematical Bioeconomics: The Optimal Management of
Renewable Resources, 2nd Ed, Wiley-Interscience, New York, ISBN 0-471-75152-9
Acknowledgment The work described in this article was supported by the Division of
Undergraduate Education of the National Science Foundation under grants DUE-0310123
and DUE-0442160. However, the views expressed are those of the author and do not
necessarily reflect those of the Foundation.
Abstract The exponential growth model and the logistic model typically introduced in
the mathematics curriculum presume that a population grows exclusively. In reality,
species can also die out and more sophisticated models that take the possibility of
extinction into account are needed. In this article, two extensions of the logistic model
are considered, one known as the logistic model with Allee effect, which is a cubic model,
and the other a further generalization, which is a quartic model. The approach used is
designed to illustrate some techniques of model-building.