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Extra info for EVOLUTION AND THELEVELS OF SELECTION
Example text
So causal decomposition is sometimes but not always possible. 3) in making these remarks. ¹⁶ Others include migration, drift, and transmission bias. This raises the question: is causal decomposition possible in this case? Can the total evolutionary change be divided into distinct components, each corresponding to a different causal factor? Biologists generally speak as if this is possible, for example, when they ask how strong selection in favour of a gene must be to prevent its elimination by drift, or how much migration is necessary to eliminate genetic variation between demes, or whether group selection in favour of a trait is strong enough to counter individual selection against it.
Prior to this stage the groups are merely loose collections of lower-level entities, not genuine evolutionary individuals. If something like this is correct, then the evolution of new levels in the hierarchy cannot be regarded as entirely prior to the evolution of adaptations at those levels. For what converts the group into a true biological unit is precisely the evolution of a special sort of group-level adaptation (cf. Frank 1995b; Szathm´ary and Wolpert 2003). I return to this issue in detail in Chapter 8, when I look at the application of multi-level selection theory to the ‘major transitions’ in evolution.
Suppose we are interested in the proportion of blue entities in our population, for some reason. We then define zi = 1 if the ith entity is blue, zi = 0 otherwise. Obviously, z then equals the proportion of blue entities in the P-population, and z the change in this proportion between the P- and O-populations. So Price’s equation applies as usual. Similarly, z could be defined as the frequency of a particular allele at a given locus in an organism (= 1, 1/2, or 0 for diploids); z would then equal the overall frequency of the allele in the population, and z the ¹² Indeed, the entities in the P- and O-populations do not need to be related as parents and offspring at all; as Price (1972) pointed out, his equation requires only an abstract correspondence between the two sets of entities.