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Course Notes for Calculus I

Section 9.3 Two Population Differential Equations

Subsection 9.3.1 Percentage Growth

In Section 3.1, I talked about percetage growth for populations. Then, in Section 3.2, I argued that I could translate percentage growth into a differential equation. If a population \(p(t)\) had a growth rate of \(c\) percent, that was equivalent to the equation
\begin{equation*} \frac{dp}{dt} = \frac{c}{100} p(t) \text{.} \end{equation*}
I don’t need to state this as percentage. If \(\alpha\) is the fraction of the population that is added or subtracted in one unit of time, the equation is
\begin{equation*} \frac{dp}{dt} = \alpha p(t) \text{.} \end{equation*}
In the history of population models, this is often called the Malthusian model of population growth. It is one of the very first pieces of mathematical biology. In the original model that Malthus produced, the constant \(\alpha\) had two (or sometimes three) pieces. There is a positive birth rate \(b\text{,}\) a negative death rate \(d\text{,}\) and possibly a migration rate \(m\) which could be positive or negative. The total growth rate is
\begin{equation*} \alpha = b - d + m \text{.} \end{equation*}
I argued, simply from regression, that the solution to the Malthusian model should be exponential. Now I’d like to prove that. The Mathusian model is a separable equation, so I can solve is using the techniques of the previous section.
\begin{equation*} \frac{dp}{dt} = \alpha p(t) \end{equation*}
In the notation of the previous section, \(f(t) = \alpha\) and \(g(p) = p\text{.}\) After seperateion, the integral become
\begin{equation*} \int \frac{1}{p} dp = \int \alpha dt \text{.} \end{equation*}
Then I can integrate each side. The integral of \(\frac{1}{p}\) is \(\ln |p|\text{.}\) Howwever, since I can assume the population function must be positive, I can drop the absolute value for an integral of \(\ln |p|\text{.}\) The integral of a constant on the right (in the variable \(t\) indicated by the \(dt\)) is \(\alpha t\text{.}\) I need a constant of integration, which I will call \(c\text{.}\)
\begin{equation*} \ln p = \alpha t + c \end{equation*}
Then I solve for \(p\text{.}\)
\begin{equation*} p(t) = e^{\alpha t + c} = e^c e^{\alpha t} \end{equation*}
If \(t = 0\text{,}\) this is \(p(0) = e^c e^0 = e^c\text{,}\) so \(e^c\) is the starting population. I’ll write this just as \(e^c = p_0\text{.}\) The solution to the percentege growth equation (or the Malthusian model) is an exponential function.
\begin{equation*} p(t) = p_0 e^{\alpha t} \end{equation*}
Being able to actually solve the DE justifies the previous connection I made between exponential functions and percentage growth. This is an ideal situation in applied mathematics: I want to observe a growth pattern, write it down in a differential equation, and actually solve that differential equation to get a function that describese the growth.
The Malthusian model is the starting point for population dymanics. A population with ample resources, space to grow, and no other limiting factors (such as predators, disease, disasters, etc.) will show percentage growth (since natural processes like birth and death rates are fractions of the totals). However, this is just the starting point. Most populations don’t have ample resources, space to grow, and freedom from other limiting factors. In Section 5.3, I already introduced the first adjustment to population dynamics: logistic growth.

Subsection 9.3.2 Logistic Growth

Logistic growth starts like exponential growth, but slows down as the population approachs a certain value, \(K\text{,}\) called the carrying capacity. This measure the effect of a population running out of excess room or resources. Essentially, the population fills its environment, or at least its niche in that environemnt, and levels off to a steady and consistent value.
Instead of adjusting the function, I want to adjust the Malthusian differential equation to model this carrying capacity. The basic model is
\begin{equation*} \frac{dp}{dt} = \alpha p\text{.} \end{equation*}
Now I want the rate of change to start out like this, but decrease as I get close to the carrying capacity. If the population is exactly at its carrying capacity, I want the growth rate to be zero. I can accomplish this by multiplying the right side by \(\left( 1 - \frac{p}{K} \right)\text{.}\) When the population is small (at least compared to \(K\)), this is almost \(1\text{,}\) so multiplication by this term has little effect. When the population is close to \(K\text{,}\) the fraction \(\frac{p}{K}\) is close to \(1\text{,}\) so the difference in brackets becomes close to \(0\text{.}\) Multiplying by something close to zero makes the growth rate shrink to a small value. This is precisely what I want: as the population approaches \(K\text{,}\) the growth rate gets smaller and smaller. Finally, if the population starts above \(K\text{,}\) this term is negative and makes the population decrease back down to this value, \(K\text{.}\)
If I include this new adjustment to the Malthusian model, I get the logistic equaiton.
\begin{equation*} \frac{dp}{dt} = \alpha p \left( 1 - \frac{p}{K} \right) \end{equation*}
I can solve this as a seperable equation as well, using the initial condition \(p(0) = p_0\) for some constant initial popluation \(p_0\text{.}\) The steps in the length process are below under as an example; they are they for your interest if you wish, but right now, I’m focused just on the result.
\begin{equation*} p(t) = \frac{Kp_0e^{\alpha t}}{ p_0e^{\alpha t} + K - p_0} = \frac{Kp_0e^{\alpha t}}{ p_0(e^{\alpha t}- 1) + K} \end{equation*}
This is ths logistic growth as it was first introduced in Section 5.3. Now it is the solution a particular differential equation.

Example 9.3.1.

Here are the steps to solve the logistic equation as a seperable differential equation.
\begin{equation*} \frac{dp}{dt} = \alpha p \left( 1 - \frac{p}{K} \right) \end{equation*}
I take \(f(t) = \alpha\) and \(g(p) = p \left( 1 - \frac{p}{K} \right)\) and use the separable technique from the previous section.
\begin{equation*} \int \frac{1}{p \left( 1 - \frac{p}{K} \right)} dp = \int \alpha dt \end{equation*}
I can simplify the nested fraction on the left by multiplying by \(K\text{.}\)
\begin{equation*} \int \frac{K}{p(K-p)} dp = \int \alpha dt \end{equation*}
Unfortunately, the left integral uses a technique that I won’t cover in this course (it is covered in Calculus II). Using a technique called partial fracitons, the left side can be split up this way.
\begin{equation*} \frac{K}{p(K-p)} = \frac{1}{p} + \frac{1}{K-p} \end{equation*}
Using linearity, this turns the left side integral into two integrals.
\begin{equation*} \int \frac{1}{p} dp + \int \frac{1}{K-p} dp = \int \alpha dt \end{equation*}
Now I can do both of these integrals. The second integral on the left uses a substitution \(u = K-p\text{,}\) for which I won’t show the steps here.
\begin{equation*} \ln p - \ln (K-p) = \alpha t + c \end{equation*}
Then I want to try to solve for \(p\text{.}\) I can first use a property of the logarithm.
\begin{equation*} \ln \frac{p}{K-p} = \alpha t + c \end{equation*}
Then I can apply the exponential to both sides.
\begin{equation*} \frac{p}{K-p} = e^{\alpha t + c} = e^c e^{\alpha t} \end{equation*}
I’ll write \(e^c = b\text{,}\) simply to have an easier way to deal with this constant.
\begin{equation*} \frac{p}{K-p} = be^{\alpha t} \end{equation*}
I’ll then take reciprocals of both sides.
\begin{equation*} \frac{K-p}{p} = \frac{1}{be^{\alpha t}} \end{equation*}
This lets me simplify the left side.
\begin{equation*} \frac{K}{p} - 1 = \frac{1}{be^{\alpha t}} \end{equation*}
Then there is only one \(p\) term, which I can isolate.
\begin{gather*} \frac{K}{p} = \frac{1}{be^{\alpha t}} + 1 = \frac{be^{\alpha t} + 1}{be^{\alpha t}}\\ \frac{p}{K} = \frac{b e^{\alpha t}}{be^{\alpha t} +1}\\ p = \frac{Kbe^{\alpha t}}{be^{\alpha t} +1} \end{gather*}
This is the logistic growth function. However, this is not precisely the form that I gave for it in Section 5.3. The reason the forms differ is that this constant \(b\) is not the starting population. If I use an initial condition \(p(0) = p_0\) in the solution, I get this expression.
\begin{equation*} p_0 = \frac{Kbe^{0}}{be^{0} +1} = \frac{Kb}{b+1} \end{equation*}
Skipping the algebra steps, I solve for \(b\text{.}\)
\begin{equation*} b = \frac{p_0}{K-p_0} \end{equation*}
I can put this into the expression.
\begin{equation*} p(t) = \frac{K\left( \frac{p_0}{K-p_0} \right) e^{\alpha t}}{ \left( \frac{p_0}{K - p_0} \right) e^{\alpha t} +1} \end{equation*}
Finally, I simplify by multiplying numerator and denominator by \((K-p_0)\)
\begin{equation*} p(t) = \frac{Kp_0e^{\alpha t}}{ p_0e^{\alpha t} + K - p_0} = \frac{Kp_0e^{\alpha t}}{ p_0(e^{\alpha t}- 1) + K} \end{equation*}
In summary, I was able to turn the two most important basic models of population into differential equations. Using the method of separable equations, I was able to actually solve the DEs and produce the functions which I claimed were the solutions in Section 3.1. I have justification for those functions, since I have now produced them from the DE instead of pulling them out of thin air.