---
OA_place: publisher
OA_type: hybrid
PlanS_conform: '1'
_id: '22998'
abstract:
- lang: eng
  text: Balancing selection, a form of selection that maintains genetic diversity,
    is difficult to detect, and the importance of balancing selection for the maintenance
    of genetic variation may be larger than often assumed. We model the possibility
    that the diversity-promoting effects of balancing selection extend to other loci
    that show sign epistasis with a locus under balancing selection. Rather than focusing
    on overdominance, as was done in previous efforts, we explore the effects of negative
    frequency dependence and show that this has important effects on the conditions
    under which the diversity-promoting effect of epistasis can occur in diploids.
    Our results show that not only recombination rate but also the dominance of sign
    epistasis are key parameters that determine the maintenance of polymorphism beyond
    the locus under direct balancing selection. We suggest that the effect we explore
    may play a significant role, especially when balancing selection acts on major
    effect loci.
acknowledgement: This work was funded by grants from the Swedish Research Council
  (2023-03730 to G.A.) and the DOC fellowship from the Austrian Academy of Science
  (26293 to K.K.). We thank L. Rowe for valuable discussions.
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Kseniia
  full_name: Khudiakova, Kseniia
  id: 4E6DC800-AE37-11E9-AC72-31CAE5697425
  last_name: Khudiakova
  orcid: 0000-0002-6246-1465
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
- first_name: Göran
  full_name: Arnqvist, Göran
  last_name: Arnqvist
biorxivid: 1
citation:
  ama: Khudiakova K, Barton NH, Arnqvist G. Sign epistasis extends the effects of
    balancing selection on genetic diversity. <i>Theoretical Population Biology</i>.
    2026;172:48-55. doi:<a href="https://doi.org/10.1016/j.tpb.2026.09.001">10.1016/j.tpb.2026.09.001</a>
  apa: Khudiakova, K., Barton, N. H., &#38; Arnqvist, G. (2026). Sign epistasis extends
    the effects of balancing selection on genetic diversity. <i>Theoretical Population
    Biology</i>. <a href="https://doi.org/10.1016/j.tpb.2026.09.001">https://doi.org/10.1016/j.tpb.2026.09.001</a>
  chicago: Khudiakova, Kseniia, Nicholas H Barton, and Göran Arnqvist. “Sign Epistasis
    Extends the Effects of Balancing Selection on Genetic Diversity.” <i>Theoretical
    Population Biology</i>, 2026. <a href="https://doi.org/10.1016/j.tpb.2026.09.001">https://doi.org/10.1016/j.tpb.2026.09.001</a>.
  ieee: K. Khudiakova, N. H. Barton, and G. Arnqvist, “Sign epistasis extends the
    effects of balancing selection on genetic diversity,” <i>Theoretical Population
    Biology</i>, vol. 172. pp. 48–55, 2026.
  ista: Khudiakova K, Barton NH, Arnqvist G. 2026. Sign epistasis extends the effects
    of balancing selection on genetic diversity. Theoretical Population Biology. 172,
    48–55.
  mla: Khudiakova, Kseniia, et al. “Sign Epistasis Extends the Effects of Balancing
    Selection on Genetic Diversity.” <i>Theoretical Population Biology</i>, vol. 172,
    2026, pp. 48–55, doi:<a href="https://doi.org/10.1016/j.tpb.2026.09.001">10.1016/j.tpb.2026.09.001</a>.
  short: K. Khudiakova, N.H. Barton, G. Arnqvist, Theoretical Population Biology 172
    (2026) 48–55.
corr_author: '1'
das_tickbox: '1'
dataavailabilitystatement: The Python code used for the numerical analyses is available
  online at https://github.com/khudyakovaks/Sign-epistasis-extends-the-effects-of-balancing-selection-on-genetic-diversity.
date_created: 2026-09-27T22:01:51Z
date_published: 2026-09-17T00:00:00Z
date_updated: 2026-10-07T06:38:02Z
day: '17'
ddc:
- '500'
- '570'
department:
- _id: JaMa
- _id: GradSch
- _id: NiBa
doi: 10.1016/j.tpb.2026.09.001
external_id:
  biorxivid:
  - 10.1101/2025.04.09.647826
  pmid:
  - '42722132'
fulldoi: https://doi.org/10.1016/j.tpb.2026.09.001
has_accepted_license: '1'
intvolume: '       172'
keyword:
- Sign epistasis
- Balancing selection
- Within-population genetic variation
language:
- iso: eng
license: https://creativecommons.org/licenses/by/4.0/
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1016/j.tpb.2026.09.001
month: '09'
oa: 1
oa_version: Published Version
page: 48-55
pmid: 1
project:
- _id: 34d33d68-11ca-11ed-8bc3-ec13763c0ca8
  grant_number: '26293'
  name: The impact of deleterious mutations on small populations
publication: Theoretical Population Biology
publication_identifier:
  eissn:
  - 1096-0325
  issn:
  - 0040-5809
publication_status: epub_ahead
quality_controlled: '1'
related_material:
  record:
  - id: '21968'
    relation: earlier_version
    status: public
researchdata_availability: no
scopus_import: '1'
status: public
supplementarymaterial: yes
title: Sign epistasis extends the effects of balancing selection on genetic diversity
tmp:
  image: /images/cc_by.png
  legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
  name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
  short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 172
year: '2026'
...