Exaptation—a missing term in the science of form

Gould, Stephen Jay; Vrba, Elisabeth S.

doi:10.1017/s0094837300004310

Cited by 5,136Philosophy and History of ScienceEvolution and Genetic DynamicsEvolution and Paleontology Studies

Exaptation—a missing term in the science of form

Gould, Stephen Jay, Vrba, Elisabeth S.·1982·Paleobiology

DOI 10.1017/s0094837300004310

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Abstract

Adaptation has been defined and recognized by two different criteria: historical genesis (features built by natural selection for their present role) and current utility (features now enhancing fitness no matter how they arose). Biologists have often failed to recognize the potential confusion between these different definitions because we have tended to view natural selection as so dominant among evolutionary mechanisms that historical process and current product become one. Yet if many features of organisms are non-adapted, but available for useful cooptation in descendants, then an important concept has no name in our lexicon (and unnamed ideas generally remain unconsidered): features that now enhance fitness but were not built by natural selection for their current role. We propose that such features be called exaptations and that adaptation be restricted, as Darwin suggested, to features built by selection for their current role. We present several examples of exaptation, indicating where a failure to conceptualize such an idea limited the range of hypotheses previously available. We explore several consequences of exaptation and propose a terminological solution to the problem of preadaptation.

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Exaptation is a shift in the function of traits wherein certain features evolve by natural selection for one function and later are co-opted to perform a different one.
“a well-known feature in the evolution of complex systems is the shift in the function of traits, called exaptation (Gould and Vrba 1982), wherein certain features evolve by natural selection for one function and later are co-opted to perform a different one.”
From Mitochondrial retrograde control of transcription evolves with respiratory stress, metabolic adaptation, and virulence in budding yeasts (2026) · cites this paper as Gould and Vrba 1982

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Authors: Gould, S. J., & Vrba, E. S.
Affiliations: Harvard University (US)
Ditsong National Museum of Natural History (ZA)
ORCID: Gould, Stephen Jay
Venue: Paleobiology
Publisher: Cambridge University Press (CUP)
Published: 1982
Volume: 8
Issue: 1
Pages: 4-15
ISSN: 0094-8373 (ISSN-L)
1938-5331
DOI: 10.1017/s0094837300004310
OpenAlex: W2106533942
Cited by: 5,136
References: 1
Citations by year: 2026: 196 · 2025: 169 · 2024: 162 · 2023: 197 · 2022: 223 · 2021: 280 · 2020: 189 · 2019: 207 · 2018: 187 · 2017: 163 · 2016: 208 · 2015: 171 · 2014: 171 · 2013: 194 · 2012: 145
FWCI: 283.14
Type: journal article
Language: en
Open access: closed
Licence: www.cambridge.org/core/terms (unspecified, 12456 d)
Locations: Paleobiology (publishedVersion)
Topics: Social Sciences › Arts and Humanities › History and Philosophy of Science › Philosophy and History of Science
Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Evolution and Genetic Dynamics
Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
Keywords: exaptation, adaptation, preadaptation, natural selection, cooption, Charles Darwin, evolutionary processes, form and function

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Exaptation—a missing term in the science of form

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