Molecular Evolution: Nearly Neutral Theory

Abstract

Nearly neutral theory is an extension of the neutral theory and contends that the borderline mutations, whose effects lie between the selected and the neutral classes, are important at the molecular level. Under the strict neutral theory, the evolutionary rate is equal to the neutral mutation rate. Under the near‐neutrality, the situation is not so simple and the most significant difference between the neutral and the nearly neutral theories is that the latter predicts a negative correlation between evolutionary rate and species population size. The nearly neutral theory also predicts abundant rare alleles in the population as compared with strict neutrality. Genome‐wide data on protein evolution are mostly in accord with the nearly neutral theory. Genetic regulatory systems are highly complex. The near‐neutrality concept may be extended to the evolution of such systems, where epigenetics and robustness are important for gene expression and many mutations are weakly selected.

Key Concepts:

  • The emphasis of significance of weak selection in evolution distinguishes the nearly neutral theory from the neutral theory.

  • The nearly neutral theory contends that the interplay of drift and weak selection is important and predicts that evolution is more rapid in small populations than in large populations.

  • Many observed patterns of protein evolution by measuring synonymous and nonsynonymous divergences are in accord with the nearly neutral theory.

  • Observed molecular polymorphisms within a population often show abundance of rare alleles, in accord with the prediction of the nearly neutral theory.

  • Numerous complex systems work together in living cells such as those in chromatin modelling/remodelling and in various signalling pathways. Interplay of drift and weak selection is important for evolution of such complex systems.

Keywords: slightly deleterious mutations; drift; effectiveness of selection; interplay of drift and selection for evolution of complex systems

Figure 1.

The classification of new mutants under the selection, neutral and nearly neutral theories. Note that while most selected mutants are deleterious, the group also includes advantageous mutants. The nearly neutral class also contains slightly deleterious and weakly advantageous mutations.

Figure 2.

Fixation probability of a mutant in a finite population as a function of 2Ns. p is the initial frequency of the mutant. The region of 2Ns<0 is that of slightly deleterious mutations.

Figure 3.

A scenario to show the meaning of near‐neutrality in relation to epigenetics and robustness that connect phenotypes with genotypes. Reproduced with permission from Ohta, .

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Further Reading

Akashi H, Osada N and Ohta T (2012) Weak selection and protein evolution. Genetics 192: 15–31.

Eyre‐Walker A and Keightley PD (2007) The distribution of fitness effects of new mutations. Nature Reviews Genetics 8: 610–618.

Gillespie JH (1991) The Causes of Molecular Evolution. Oxford: Oxford University Press.

Hartl DL and Clark A (2007) Principles of Population Genetics, 4th edn. Sunderland, MA: Sinauer.

Lynch M and Cornery JS (2003) The origins of genome complexity. Science 302: 1401–1404.

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How to Cite close
Ohta, Tomoko(Feb 2013) Molecular Evolution: Nearly Neutral Theory. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001801.pub4]