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Evolution__3rd_Edition

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304 PART 3 / Adaptation and Natural Selection<br />

(a)<br />

Day 1<br />

Day 37<br />

Breeds for<br />

37 days<br />

48 new colonies<br />

(16 beetles each)<br />

(b)<br />

Mean number of adults<br />

300<br />

250<br />

200<br />

150<br />

100<br />

50<br />

48 colonies<br />

(16 beetles in a colony)<br />

0<br />

0 1 2 3 4 5 6 7 8 9<br />

Generations<br />

Group-selected beetles evolve<br />

lower fecundity<br />

High<br />

Control<br />

Low<br />

Figure 11.2<br />

Wade’s experiment with Tribolium beetles. (a) Experimental<br />

design: 48 colonies were bred for 37 days, and then a new round<br />

of colonies was formed from the colonies that had grown to a<br />

low (or high) population density. (b) Results showing population<br />

densities in lines selected for high or low population densities and<br />

in unselected controls. The results are the means for 48 colonies.<br />

Redrawn, by permission of the publisher, from Wade (1976).<br />

Not surprisingly, the population density in the “low” lines decreased relative to the<br />

“high” lines (Figure 11.2b). The decrease in the low lines is due to group selection. Presumably,<br />

within the 37 days of any one cycle, the beetle types with high fecundity were<br />

increasing within each colony relative to the less fecund beetles. However, between<br />

cycles, Wade’s group selection for low fecundity more than out-weighed the individual<br />

selection and average fecundity declined. The group selection was strong enough to work.<br />

In a way, the group selective structure of the experiment is superfluous. We could<br />

simply breed from beetles with lower fecundity. Artificial selection of this kind would<br />

reduce beetle fecundity without their being kept in groups for 37 days. But Wade’s purpose<br />

was to illustrate group, not artificial, selection and his experiment does so. It has<br />

alternating rounds of individual and group selection and the experimental group selection<br />

is strong enough to produce the effect that Wynne-Edwards thought to be common<br />

in nature. Box 11.1 shows how the group selection design of Wade’s experiment<br />

has had a practical application.<br />

The fact that group selection can be implemented in an experiment does not mean<br />

that group selection is important in nature. Biologists doubt group selection for theoretical<br />

reasons, and because of the kinds of adaptations seen in nature. The experiments<br />

are instructive, however. They show what group selection means, and how individual<br />

selection can decrease the efficiency of a group. Muir’s experiment (Box 11.1) also has a<br />

commercial interest.<br />

..

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