Population Dynamics 2 Geometric and Exponential Growth Model Revision This section will start with a revision of the

previous section. Question 1: Do you remember the general model of population growth? Answer: $ N_{t+1} = Nt + B + I - D - E $ (Equation 1)

Question 2: Can you define the parameters: B, D, I, E? Answer: B = the number of births from t to t+1 (birth rate or natality), D = the number of deaths from t to t+1 (death rate or mortality), I = the number of immigrants from t to t+1 (immigration rate), and E = the number of emigrants from t to t+1 (emigration rate). Question 3: Do you remember how Equation 1 changes when we describe the population dynamics of a closed population? Why the equation assumes this form? Answer: $ N_{t+1} = Nt + B - D $ (Equation 2)

In a closed population there are no Immigration (I) and Emigration (E), thus I=E=0 For simplicity, in this section, we will use only the population dynamics model of a closed population (Equation 2).

Geometric Growth Now, as discussed in the Population Increase or Decline section, the per capita rate is a rate per individual; so the per capita birth rate (bt) is the number of births per individual in the population per unit time, and the per capita death rate (dt) is the number of deaths per individual in the population per unit time. That is, per capita birth and death rates are each individuals probability of giving birth or dying in a given unit of time. Therefore, to calculate per capita rates, we divide Bt the total number of births in the interval from time t to time t + 1 and Dt the total number of births in the same interval by the population size. Thus, bt= Bt/Nt and dt = Dt/Nt
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Conversely, Bt= btNt and Dt= dtNt (Equation 3)

Now we can rewrite our model (Equation 2) in terms of per capita rates: Nt+1 = Nt+ btNt dtNt (Equation 4)

Let us assume that average number of births per unit time per individual in the population and the average risk of dying per unit time remain unchanged over some period. What will happen to population size? Because we assume constant per capita birth and death rates, we can make one further, minor modification to our equation by leaving off the time subscripts on b and d: Because per capita birth and death rates do not change in response to the size (or density) of the population, this model is said to be density-independent. We can further simplify Equation 4 by factoring Nt out of the birth and death terms: Nt+1= Nt + (b d)Nt (Equation 5)

The term (b d) is so important in population biology that it is given its own symbol, R. Thus R = b d, and is called the geometric rate of increase. Substituting R for (b d) gives us Nt+1= Nt+ RNt (Equation 6)

To further define R, we can calculate the rate of change in population size, DeltaNt, by subtracting Nt from both sides of Equation 6: DeltaNt= Nt+1 Nt = RNt Because DeltaNt= Nt+1 Nt , we can simply write DeltaNt = RNt (Equation 7)

In other words, the rate of change in population size is proportional to the population size, and the constant of proportionality is R. Question: Do you remember what are the per capita population parameters and how to calculate them? Answer: Per capita birth rate (b), (also called nativity rate), is the number of births per individual per time step. It is calculated as the number of births (B) divided by the size of the reproductive population (N): b = B/N or bN = B

Per capita death rate, (also called mortality rate), is the number of deaths per individual per time, calculated as the number of deaths, divided by the step: d = D/N or dN = D Now, let us get back to our discussion of the model of the geometric population growth.

As discussed, the rate of change in population size is proportional to the population size, and the constant of proportionality is R, so DeltaNt = RNt (Equation 7)

We can convert Equation 7 to per capita rate of change in population size if we divide both sides of the equation by Nt : DeltaNt/ Nt =R In other words, the parameter R represents the (discrete-time) per capita rate of change in the size of the population. Moving on, we can simplify Equation 6 (Nt+ 1 = Nt + RN) even further by factoring Nt out of the terms on the right-hand side, to get Nt+1 = (1 + R)Nt The quantity (1 + R) is often given its own symbol, (lambda), and its own name: the finite rate of increase. Substituting , we can write Nt+ 1 = Nt (Equation 8)

If we divide both sides of Equation 8 by Nt, we get: N t +1/ Nt = (Equation 9)

In other words, is the ratio of the population size at one time to its size one time-unit earlier. We can calculate from population counts at successive times, even if we do not know per capita rates of birth and death. Furthermore when, > 1, exponential increase = 1, no changestationary population < 1, exponential decline

Excersize: If the population size of a pack of wolves in a certain area is 150 in year 1 and the size is 200 in the year 2, what is the value of lambda? Is the population experiencing an exponential increase, an exponential decline, or is it a stationary population? Answer: From Equation 6: N t +1/ Nt = Thus, 200/150=1.33

Since (1.33) > 1, the population of wolves is experiencing an exponential increase. Excersize: For the same population of wolves, which numbers 150 individuals in year 1, what would be the population size in year 2, if = 1.20? Answer: N t +1/ Nt = , and Nt=150, = 1.20 Then N 2/150=1.20 N 2/150=1.20 N 2=150x1.20 N 2=180 wolves Thus in year 2 the population would increase to 180 wolves, that is - by 30 wolves. Equation 6 to 9 demonstrated how to calculate the population size one time unit step into the future. What if you wanted to know how big the population will be at some distant future time? You could carry out the one-time-step calculations many times, until you arrived at the desired answer, or you could use a shortcut. Let us start with Equation 8: Nt+1= Nt Starting at time 0, we can carry this calculation through a few times to calculate population sizes at time 1, time 2, and time 3. The population size at time 0 can be written N0. Thus the populations at times 1, 2, and 3 would be: N1= N0 N2= N1= ( N0) N3= N2= ( ( N0) ) This development represents a pattern: Population size at time 1 is 1N0, at time 2 it is 2N0, and at time 3 is 3N0 Thus, in general, we can write Nt= tN0 Equation (10)

Exercise: A pack of wolves is described by a size N0 = 100 and = 1.15. What would be the total population size in t = 10 years

Solution: Nt= tN0 N10 = (1.1510)100 = 405 The total population size in 10 years will be 405 wolves. Now, let us return to our discussion. An important application of this model of population growth is to use Equation 10 to calculate doubling time ( tdouble) - that is, the time required for the population to double in size. If we plug the doubling time into Equation 10, we determine that: Nt(double)= t(double)N0 We can derive doubling time by exploiting the fact that the population at time tdouble is, by definition, twice the population at time 0: Ntdouble= 2N0 2N0= t(double)N0 If we divide both sides by N0, we get 2= t(double) Taking the logarithm of both sides gives us: ln2 =t(double)ln() Dividing both sides by ln, we get: ln(2)/ ln()= t(double) (Equation 11)

What does this mean? Suppose R = 0.2 individuals/individual/year. Therefore, = 1 + R = 1.2. This implies that the population increases by 20% per year, which doesnt seem like a too big increase. But, if you plug this value of into Equation 11, youll find that the population doubles in only about 3.80 years! Since a population that grows in discrete intervals of a year can double in a non-integer number of years, this calculation also means that the population will not quite double in 3 years, and will more than double in 4 years.