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one that returns to the original equilibrium point after being perturbed. An unstable equilibrium is one that will not return after being perturbed but, rather, continues to move away from the equilibrium point. As we mentioned in the last chapter, the Hardy-Weinberg equilibrium is a neutral equilibrium: It remains at the allelic frequency it moved to when perturbed. Stable, unstable, and neutral equilibrium points can be visualized as marbles in the bottom of a concave surface (stable), on the top of a convex surface (unstable), or on a level plane (neutral; g. 20.1). Although more sophisticated mathematical formulas exist for determining whether an equilibrium is stable, unstable, or neutral, we will use graphical analysis for this purpose. Figure 20.2 introduces the process of graphical analysis, which provides an understanding of the dynamics of an event or process by representing the event in graphical form. In gure 20.2, we have graphed equation 20.3, the q equation of mutational dynamics.The ordinate, or y-axis, is q, the change in allelic frequency.The abscissa, or x-axis, is q, or allelic frequency.The diagonal line is the q equation, the relationship between q and q. Note that q can be positive (q is increasing) or negative (q is decreasing), whereas q is always positive (0 1.0). Graphical analysis can provide insights into the dynamics of many processes in population genetics. The diagonal line in gure 20.2 crosses the q 0 ^ line at the equilibrium value (q ) of 0.167. This line also shows us the changes in allelic frequency that occur in a population not at the equilibrium point. We will look

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at two examples of populations under the in uence of mutation pressure, but not at equilibrium: one at q 0.1 (below equilibrium) and one at q 0.9 (above equilibrium). If we substitute q 0.1 into equation 20.3, we get a q value of 4 10 6. If we substitute q 0.9 into the equation, we get a q value of 4.4 10 5. In other words, when the population is below equilibrium, q increases ( q 4 10 6 ); if the population is above equilibrium, q decreases ( q 4.4 10 5). We can read these same conclusions directly from the graph in gure 20.2. We can see that the mutational equilibrium is a stable one. Any population whose allelic frequency is not at the equilibrium value tends to return to that equilibrium value. A shortcoming of this model is that it provides no obvious information revealing the time frame for reaching equilibrium. To derive the equations needed to determine this parameter is beyond our scope. (We could use computer simulation or integrate equation 20.3 with respect to time.) In a large population, any great change in allelic frequency caused by mutation pressure alone takes an extremely long time. Most mutation rates are on

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Graphical analysis of mutational equilibrium. The graph of the mutational q equation shows that when the population is perturbed from the equilibrium point (q 0.167), it returns to that equilibrium point. At q values above equilibrium, change is negative, tending to return the population to equilibrium. At q values below equilibrium, change is positive, also tending to return the population to equilibrium.

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the order of 10 5, and equation 20.3 shows that change will be very slow with values of this magnitude. For example, if 10 5, 10 6, and p q 0.5, q (0.5 10 5 ) (0.5 10 6 ) 4.5 10 6, or 0.0000045. It usually takes thousands of generations to get near equilibrium, which is approached asymptotically. As you can see from the low values of mutation rates, it would usually be nearly impossible to detect perturbations to the Hardy-Weinberg equilibrium by mutation in any one generation. The mutation rate can, however, determine the eventual allelic frequencies at equilibrium if no other factors act to perturb the gradual changes that mutation rates cause. Mutation can also affect nal allelic frequencies when it restores alleles that natural selection is removing, a situation we will discuss at the end of the chapter. More important, mutation provides the alternative alleles that natural selection acts upon.

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