Summary
Moderately diverged small sets of nonallelic genes that control one type of protein, such as genes for isozymes, are usually considered to be present in order to permit adaptations of protein function to local conditions in organismal time and space. However some irregularities, as well as some regularities, in the distribution of lactate dehydrogenase isozymes seem difficult to reconcile with this view. A complementary interpretation is based on selection for gene regulation. Irrespective of the relative importance of transcriptional control and of control through processing of the transcripts, regulatory patterns in higher organisms may be explored on the basis of formal relationships between molecular components interacting within and between “controller nodes”. It is shown that an analysis in terms of controller node stability states may account for the variations in regulation of isozyme synthesis. Adaptive changes in regulatory sequences may insure the stability of the synthesis of a given type of protein in the face of developmental, as well as evolutionary, alterations in patterns of gene activity, correlated with the evolution of cell differentiation. It is proposed that gene duplication, beyond the addition of novel functions to the established ones, plays a further and rather opposite role. This role is to maintain theconstancy of established protein functions in the presence of developmental and evolutionary change. To this effect the duplication of functional units of gene action, rather than merely of coding sequences, is likely to be required.
In a certain number of cases, higher organisms can apparently dispense with a given moderately diverged set of genes and use a single gene instead. This fact is, to some extent, at variance with both the protein-functional and regulatory interpretations of the existence of the set. The relevance of the concept of genetic sufficiency is suggested as a solution, and possibilities offered by this concept for quantitative investigations in population genetics are outlined in an appendix. Increases in fitness occur automatically and perpetually regardless of the fact that genetic sufficiency often has already been reached. This process is characterized as the fitness treadmill. The reason why higher organisms evolve from lower forms probably is that sufficiency in adaptedness in no way prevents the fitness treadmill from continuing to turn. Tolerance towards gene loss does not imply that the dispensable genes have not continuously been selected for. It is proposed that increases and decreases during evolution in haploid DNA content per cell occur according to whether the fitness treadmill alone operates constantly, or whether frequent episodes of population bottlenecks bring genetic sufficiency thresholds to the fore. Positive selection does notper se confer biological significance upon a mutant. The frequency of biologically important selection events is expected to depend on the variability of the environment. The approximately equal evolutionary rates of amino acid replacements in a given kind of protein in both evolutionarily relatively stable and unstable environments suggest that most increases in fitness occur under conditions of continued genetic sufficiency of the wild type that is being replaced by a mutant. In relation to the potential for survival of the species and lineage, most selected mutations probably are no more, nor less, important than neutral mutations.
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Zuckerkandl, E. Multilocus enzymes, gene regulation, and genetic sufficiency. J Mol Evol 12, 57–89 (1978). https://doi.org/10.1007/BF01732545
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DOI: https://doi.org/10.1007/BF01732545