## Concepts

Natural selection is the differential reproduction of genotypes. It is measured as fitness, which is the reproductive success of a genotype compared with other genotypes in a population.

The general selection model Differential fitness among genotypes over time leads to changes in the frequencies of the genotypes, which, in turn, lead to changes in the frequencies of the alleles that make up the genotypes. We can predict the effect of natural selection on allelic frequencies by using a general selection model, which is outlined in Table 23.4. Use of this model requires knowledge of both the initial allelic frequencies and the fitness values of the genotypes. It assumes that mating is random and the only force acting on a population is natural selection.

We have defined fitness in terms of relative reproduction, but it will be easier to understand the logic behind the general selection model if we think of the fitness of the genotypes as differences in survival. It applies equally to fitnesses representing differential reproduction.

Let's apply the general selection model outlined in Table 23.4. Imagine a flock of sparrows overwintering in Rochester, New York. Assume that we can determine the genotypes for a locus that affects the ability of the birds to survive the winter; perhaps the genes at this locus determine the amount of fat that a bird accumulates before the onset of winter. For genotypes A1A1, A1A2, and A2A2, p rep resents the frequency of A1 and q represents the frequency of A2. On the first line of the table, we record the initial genotypic frequencies before selection has acted, before the onset of winter. If mating has been random (an assumption of the model), the genotypes will have the Hardy-Weinberg equilibrium frequencies of p2, 2pq, and q2. On the second row of the table, we put the fitness values of the corresponding genotypes. Some of the birds die in the winter; so here the fitness values represent the relative survival of the three genotypes. The proportion of the population represented by each genotype after selection is obtained by multiplying the initial genotypic frequency times its fitness (third row of Table 23.4). Now the genotypes are no longer in Hardy-Weinberg equilibrium.

The mean fitness (W) of the population is the sum of the proportionate contributions of the three genotypes:

The mean fitness W is the average fitness of all individuals in the population and allows the frequencies of the genotypes after selection to be obtained. In our flock of birds, these frequencies will be those of the three genotypes after the winter mortality. The frequency of a genotype after selection will be equal to its proportionate contribution divided by the mean fitness of the population (p2W11/W for genotype A1A1, 2pqW12/W for genotype A1A2, and q2W22 /W for genotype A2A2), as shown in the fourth line of Table 23.4. When the new genotypic frequencies have been calculated, the new allelic frequency of A1 (p') can be determined by using the now-familiar formula (Equation 23.4):

p' = f(A1) = f(A1A1) + 72 f(A1A2) and that of q' can be obtained by subtraction:

Method for determining changes in allelic frequency due to selection

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