Derivation of equation (2-2) on page 23

First note the errata sheet that this equation was botched by the printer.

We consider template replication to involve basically two processes, although this is a considerable oversimplification of many intermediate steps which, if modeled, do not change the basic form. First, template plus energy-rich mononucleotides (the resources in this simple system) form a template-resource complex. Second, this complex dissociates into two templates. Let denote replicator and its limiting resource (or substrate). The derivation follows the Michaelis-Menten kinetics common in the chemistry of enzyme kinetics (see, for example, a popular text in biochemistry, that of Leninger, pp. 154-155 in an old version that is on my shelves). The template functions as the "enzyme," the resources function as the substrate, and the product is then a new complementary strand plus the old template (which is unused in the reaction, although it catalyzes the formation of the complementary strand). The notation means concentration of replicator ; etc. for . We consider here only template replication with no enzyme, although the approach can be extended to enzyme mediated replication. The overall birth process is described by the following diagram:

intermediary complex:

The rate of formation of complex is .

The rate of breakdown of the complex is .

The reaction is in steady-state when the concentration of replicator-resource complex is constant or setting the rate of formation of complex to its rate of breakdown gives

,

or .

Following the Michaelis-Menten kinetics it can be seen that is the concentration of resources (mononucleotide building blocks) at which the rate of growth is half maximum. Solving this for the steady-state value of template resource complex gives

.

Since the initial rate of replication of is proportional to the concentration of complex:

.

When resources are common, the rate of replication no longer depends on resource concentration and is directly proportional to the concentration of replicators

substitute into above gives:

Upon dividing both sides of the last equation by the maximum rate of change in concentration of replicator i, , gives

or solving for

recall that on , so we have

However there is also the death process which is independent of resources. The total is then the sum of the birth process and the death process to give

.

Note that in the absence of resource competition, the growth rate of the replicator is exponential, with rate . I refer to such replicators as "Malthusian" because they obey Malthus' law of exponential increase when uninhibited by resource limitation or competition. Another derivation (Szathmary 1989) of template replication gives a square root growth law: . This can be referred to as subexponential growth, since the basic growth process is less than exponential. The reason for this is that the template will readily form a duplex with its complement and is then temporarily unavailable for replication, in other words each template interferes with its own replication more than it interferes with the replication of a competitor. The form of the replication dynamic dramatically affects the condition for selection. Malthusian growth leads to "survival of the fittest" while subexponential growth leads to a cost of commonness or "survival of anybody." More than exponential growth (as exists with protein catalyzed replication or with sexual reproduction), say , leads to a cost of rarity and "survival of the first."