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A century of theories of balancing selection

Ruzicka, Filip ; Zwoinska, Martyna K. ; Goedert, Debora LU ; Kokko, Hanna ; Li Richter, Xiang Yi ; Moodie, Iain R. LU orcid ; Nilén, Sofie LU orcid ; Olito, Colin LU ; Svensson, Erik I. LU orcid and Czuppon, Peter , et al. (2025) In Biological Reviews
Abstract

Traits that affect organismal fitness are often highly genetically variable. This genetic variation is vital for populations to adapt to their environments, but it is also surprising given that nature – after all – ‘selects’ the best genotypes at the expense of those that fall short. Explaining the extensive genetic variation of fitness-related traits is thus a longstanding puzzle in evolutionary biology, with cascading implications for ecology, conservation, and human health. Balancing selection – an umbrella term for scenarios in which natural selection maintains genetic variation – is a century-old explanation to resolve this puzzle that has gained recent momentum from genome-scale methods for detecting it. Yet evaluating whether... (More)

Traits that affect organismal fitness are often highly genetically variable. This genetic variation is vital for populations to adapt to their environments, but it is also surprising given that nature – after all – ‘selects’ the best genotypes at the expense of those that fall short. Explaining the extensive genetic variation of fitness-related traits is thus a longstanding puzzle in evolutionary biology, with cascading implications for ecology, conservation, and human health. Balancing selection – an umbrella term for scenarios in which natural selection maintains genetic variation – is a century-old explanation to resolve this puzzle that has gained recent momentum from genome-scale methods for detecting it. Yet evaluating whether balancing selection can, in fact, resolve the puzzle is challenging, given the logistical constraints of distinguishing balancing selection from alternative hypotheses and the daunting collection of theoretical models that formally underpin this debate. Here, we track the development of balancing selection theory over the last century and provide an accessible review of this rich collection of models. We first outline the range of biological scenarios that can generate balancing selection. We then examine how fundamental features of genetic systems – non-random mating between individuals, ploidy levels, genetic drift, linkage, and genetic architectures of traits – have been progressively incorporated into the theory. We end by linking these theoretical predictions to ongoing empirical efforts to understand the evolutionary processes that explain genetic variation.

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organization
publishing date
type
Contribution to journal
publication status
epub
subject
keywords
balancing selection, evolutionary theory, fitness variation, heterozygote advantage, mathematical modelling, negative frequency-dependent selection, population genetics, trade-offs
in
Biological Reviews
publisher
John Wiley & Sons Inc.
external identifiers
  • scopus:105021540183
  • pmid:41235821
ISSN
1464-7931
DOI
10.1111/brv.70103
language
English
LU publication?
yes
id
7e0ed135-1c8e-45a1-832c-24cffe808cd1
date added to LUP
2026-01-08 14:49:57
date last changed
2026-01-09 14:43:16
@article{7e0ed135-1c8e-45a1-832c-24cffe808cd1,
  abstract     = {{<p>Traits that affect organismal fitness are often highly genetically variable. This genetic variation is vital for populations to adapt to their environments, but it is also surprising given that nature – after all – ‘selects’ the best genotypes at the expense of those that fall short. Explaining the extensive genetic variation of fitness-related traits is thus a longstanding puzzle in evolutionary biology, with cascading implications for ecology, conservation, and human health. Balancing selection – an umbrella term for scenarios in which natural selection maintains genetic variation – is a century-old explanation to resolve this puzzle that has gained recent momentum from genome-scale methods for detecting it. Yet evaluating whether balancing selection can, in fact, resolve the puzzle is challenging, given the logistical constraints of distinguishing balancing selection from alternative hypotheses and the daunting collection of theoretical models that formally underpin this debate. Here, we track the development of balancing selection theory over the last century and provide an accessible review of this rich collection of models. We first outline the range of biological scenarios that can generate balancing selection. We then examine how fundamental features of genetic systems – non-random mating between individuals, ploidy levels, genetic drift, linkage, and genetic architectures of traits – have been progressively incorporated into the theory. We end by linking these theoretical predictions to ongoing empirical efforts to understand the evolutionary processes that explain genetic variation.</p>}},
  author       = {{Ruzicka, Filip and Zwoinska, Martyna K. and Goedert, Debora and Kokko, Hanna and Li Richter, Xiang Yi and Moodie, Iain R. and Nilén, Sofie and Olito, Colin and Svensson, Erik I. and Czuppon, Peter and Connallon, Tim}},
  issn         = {{1464-7931}},
  keywords     = {{balancing selection; evolutionary theory; fitness variation; heterozygote advantage; mathematical modelling; negative frequency-dependent selection; population genetics; trade-offs}},
  language     = {{eng}},
  month        = {{11}},
  publisher    = {{John Wiley & Sons Inc.}},
  series       = {{Biological Reviews}},
  title        = {{A century of theories of balancing selection}},
  url          = {{http://dx.doi.org/10.1111/brv.70103}},
  doi          = {{10.1111/brv.70103}},
  year         = {{2025}},
}