SCIENCE CHINA Life Sciences, Volume 62, Issue 4: 437-452(2019) https://doi.org/10.1007/s11427-018-9447-8

Exaptation at the molecular genetic level

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  • ReceivedNov 10, 2018
  • AcceptedDec 1, 2018
  • PublishedDec 12, 2018


The realization that body parts of animals and plants can be recruited or coopted for novel functions dates back to, or even predates the observations of Darwin. S.J. Gould and E.S. Vrba recognized a mode of evolution of characters that differs from adaptation. The umbrella term aptation was supplemented with the concept of exaptation. Unlike adaptations, which are restricted to features built by selection for their current role, exaptations are features that currently enhance fitness, even though their present role was not a result of natural selection. Exaptations can also arise from nonaptations; these are characters which had previously been evolving neutrally. All nonaptations are potential exaptations. The concept of exaptation was expanded to the molecular genetic level which aided greatly in understanding the enormous potential of neutrally evolving repetitive DNA—including transposed elements, formerly considered junk DNA—for the evolution of genes and genomes. The distinction between adaptations and exaptations is outlined in this review and examples are given. Also elaborated on is the fact that such distinctions are sometimes more difficult to determine; this is a widespread phenomenon in biology, where continua abound and clear borders between states and definitions are rare.


Apologies to those whose articles were not cited which, in part, is owed to the explosive growth of the literature in the field. The author is grateful to Stephanie Klco-Brosius for a last minute review of language.

Interest statement

The author(s) declare that they have no conflict of interest.


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  • Figure 1

    Definition of aptation, adaptation and exaptation. This figure is adapted from Table 1 in the original publication of Gould and Vrba (1982), in which the distinction of exaptation out of a character previously shaped by selection for a given function (IIa) as well as out of a neutrally evolving character is illustrated (IIb) in comparison to adaptation.

  • Figure 2

    (Color online) The continuum of aptations: fuzzy boundaries between adaptations (bottom, red area) and exaptations (top, blue area). An adaptation describes the event(s) by which point mutations or small indels modify protein or RNA coding regions, or regulatory elements, altering the corresponding gene products or the action of regulatory regions to the advantage of the carrier organism. De novo gene formation from neutrally evolving sequences and horizontal gene transfer clearly correspond to exaptation. Duplication of a gene (either by recombination or retrotransposition) can yield genes with variant or novel function, sometimes via neutrally evolving inactive stages. These are exaptations. If a gene duplicates and remains largely unaltered, for example, with the effect of a beneficial higher expression of the corresponding product, this would be similar to a point mutation in the promoter region and thus, could also be delineated as adaptation. A gene residing after duplication on an autosome, escaping from inactivation on a sex chromosome, can be placed somewhere in the no man’s land between exaptation and adaptation. Subfunctionalization of genes after duplication could also be in this nebulous area. Point mutation of a neutrally evolving sequence fortuitously generating a transcription factor (TF) binding site is closer to adaptation, whereas the recruitment of novel exons or regulatory regions out of neutrally evolving sequences (including TEs) are usually considered to be exaptations. Inactivation of a gene encoding a protein and subsequent neutral evolution or continued transcription and maintained function as non-protein coding RNA (bottom part of box on the left) could be considered a partial nonaptation with the continued potential of adaptation (of the RNA function). However, neutral evolution of the open reading frame (ORF) and subsequent revival of a protein coding region that is different from the original one clearly falls under exaptation (top part of box on the left).

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