Smith, John Maynard, and Eors Szathmáry. “The evolution of templates.” In The Major Transitions in Evolution . Oxford University Press, 1995.
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In this chapter, we discuss the origin and early evolution of genetic replication. The argument is complex, so we start with a brief outline. Section 4.2 discusses the nature of replication. We draw a distinction between simple replicators, limited hereditary replicators and indefinite hereditary replicators. Continued evolution requires indefinite hereditary replicators: it seems that such replicators depend on some form of template reproduction. In section 4.3, we point out that there is an error threshold for the accuracy of replication: for a given total quantity of genetic information–for example, for a fixed number of bases—there is an upper limit on the error rate of replication. If the error rate rises above this limit, natural selection cannot maintain the information. This leads to what we have called Eigen’s paradox. In the absence of specific enzymes, replication accuracy is low. Hence the total genome size must be small–almost certainly, less than 100 nucleotides. The genome is therefore too small to code for accurate replication machinery. There is a catch-22 situation: no enzymes without a large genome, and no large genome without enzymes. The next three sections discuss possible solutions to the paradox. Section 4.4 considers populations of replicating RNA molecules. We point out that the dynamics of replication are such as to lead to the stable coexistence of a diverse population, but we do not think that this constitutes a solution to the paradox. Section 4.5 discusses the hypercycle, a particular relationship between replicators that makes it possible for a greater total quantity of information to be maintained with a given accuracy of replication. We argue that the further evolution of hypercycles requires that they be enclosed within compartments, because otherwise they are sensitive to parasitic replicators. We also discuss, rather inconclusively, the possibility that, even in the absence of compartments, cooperation might evolve, by a processes analogous to kin selection, if the components of the hypercycle were confined to a surface. Finally, we discuss an alternative model, the stochastic corrector model. This also depends on the existence of compartments, but emphasizes the importance of stochastic effects arising if there are small numbers of each kind of molecule in a compartment. Essentially, small numbers serve to generate variation upon which selection can act.