---
OA_type: closed access
_id: '21900'
abstract:
- lang: eng
text: Individually silencing 125 fruit fly genes reveals opposing fitness effects
of mutations between females and males, as well as between germline and somatic
tissues.
article_processing_charge: No
article_type: comment
author:
- first_name: Filip
full_name: Ruzicka, Filip
id: 347955dd-57b0-11ee-9095-c28bdd368f4b
last_name: Ruzicka
citation:
ama: Ruzicka F. Reverse genetics of sexual antagonism. Nature Ecology & Evolution.
2026. doi:10.1038/s41559-026-03036-y
apa: Ruzicka, F. (2026). Reverse genetics of sexual antagonism. Nature Ecology
& Evolution. Springer Nature. https://doi.org/10.1038/s41559-026-03036-y
chicago: Ruzicka, Filip. “Reverse Genetics of Sexual Antagonism.” Nature Ecology
& Evolution. Springer Nature, 2026. https://doi.org/10.1038/s41559-026-03036-y.
ieee: F. Ruzicka, “Reverse genetics of sexual antagonism,” Nature Ecology &
Evolution. Springer Nature, 2026.
ista: Ruzicka F. 2026. Reverse genetics of sexual antagonism. Nature Ecology &
Evolution.
mla: Ruzicka, Filip. “Reverse Genetics of Sexual Antagonism.” Nature Ecology
& Evolution, Springer Nature, 2026, doi:10.1038/s41559-026-03036-y.
short: F. Ruzicka, Nature Ecology & Evolution (2026).
corr_author: '1'
date_created: 2026-05-20T14:36:45Z
date_published: 2026-05-01T00:00:00Z
date_updated: 2026-05-21T05:49:25Z
day: '01'
department:
- _id: BeVi
doi: 10.1038/s41559-026-03036-y
language:
- iso: eng
month: '05'
oa_version: None
publication: Nature Ecology & Evolution
publication_identifier:
eissn:
- 2397-334X
publication_status: epub_ahead
publisher: Springer Nature
quality_controlled: '1'
scopus_import: '1'
status: public
title: Reverse genetics of sexual antagonism
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
year: '2026'
...
---
OA_place: publisher
OA_type: hybrid
PlanS_conform: '1'
_id: '20044'
abstract:
- lang: eng
text: Genetic trade-offs—which occur when variants that are beneficial in some contexts
of natural selection are harmful in others—can influence a wide range of evolutionary
phenomena, from the maintenance of genetic variation to the evolution of aging
and sex differences. An extensive body of evolutionary theory has focused on the
consequences of such trade-offs, and recent analyses of Fisher’s geometric model
have further quantified the expected proportion of new mutations that exhibit
trade-offs. However, the theory remains silent regarding the prevalence of trade-offs
among the variants that contribute to adaptation. Here, we extend Fisher’s geometric
model to predict the prevalence of trade-offs among the adaptive mutations that
become established or fixed in a population. We consider trade-offs between sexes,
habitats, fitness components, and temporally fluctuating environments. In all
4 scenarios, trade-off alleles are consistently under-represented among established
relative to new beneficial mutations—an effect that arises from the greater susceptibility
of trade-off alleles to genetic drift. Adaptation during a population size decline
exacerbates this deficit of trade-offs among established mutations, whereas population
expansions dampen it. Consequently, threatened populations should primarily adapt
using unconditionally beneficial alleles, while invasive populations are more
prone to adaptation using variants that exhibit trade-offs.
acknowledgement: 'Support for this research came from the European Society of Evolutionary
Biology (ESEB) through a “Special Topics Network” grant. Further financial support
came from the European Research Council (ERC-2023-STG916 #101117517, to C.O.), the
Swedish Research Council (#2022-03603, to C.O.; #2020‑03123, to E.I.S.), the Research
Council of Norway (Norges forskningsråd #302619, to D.G.), the Alexander von Humboldt
Foundation and the GenEvo graduate school (to H.K.), the Foundation for Zoological
Research and the Birgitta Sintring Foundation (#S2024-0007, to M.K.Z.), a postdoctoral
fellowship from the Consejo Nacional de Humanidades, Ciencias y Tecnología (to A.N.B.),
and a H2020 Marie Skłodowska-Curie COFUND Action fellowship (#101034413, to F.R.).
We wish to express our deepest gratitude to Lotte de Vries for extensive discussion
of the project, rederiving some of our results, and providing comments on an earlier
version of the manuscript, and to the European Society of Evolutionary Biology (ESEB)
for a “Special Topics Network” grant that supported workshops that initiated this
collaboration and facilitated many new ideas and friendships. We also thank two
anonymous reviewers for their thoughtful comments and suggestions that helped us
to substantially improve upon the original version of the article.'
article_processing_charge: Yes (in subscription journal)
article_type: original
author:
- first_name: Tim
full_name: Connallon, Tim
last_name: Connallon
- first_name: Peter
full_name: Czuppon, Peter
last_name: Czuppon
- first_name: Colin
full_name: Olito, Colin
last_name: Olito
- first_name: Debora
full_name: Goedert, Debora
last_name: Goedert
- first_name: Hanna
full_name: Kokko, Hanna
last_name: Kokko
- first_name: Angela
full_name: Nava-Bolaños, Angela
last_name: Nava-Bolaños
- first_name: Sofie
full_name: Nilén, Sofie
last_name: Nilén
- first_name: Erik I
full_name: Svensson, Erik I
last_name: Svensson
- first_name: Martyna
full_name: Zwoinska, Martyna
last_name: Zwoinska
- first_name: Ludovic
full_name: Dutoit, Ludovic
last_name: Dutoit
- first_name: Filip
full_name: Ruzicka, Filip
id: 347955dd-57b0-11ee-9095-c28bdd368f4b
last_name: Ruzicka
citation:
ama: Connallon T, Czuppon P, Olito C, et al. Predicting the prevalence of genetic
trade-offs among adaptive substitutions. Evolution. 2025;79(7):1243-1255.
doi:10.1093/evolut/qpaf061
apa: Connallon, T., Czuppon, P., Olito, C., Goedert, D., Kokko, H., Nava-Bolaños,
A., … Ruzicka, F. (2025). Predicting the prevalence of genetic trade-offs among
adaptive substitutions. Evolution. Oxford University Press. https://doi.org/10.1093/evolut/qpaf061
chicago: Connallon, Tim, Peter Czuppon, Colin Olito, Debora Goedert, Hanna Kokko,
Angela Nava-Bolaños, Sofie Nilén, et al. “Predicting the Prevalence of Genetic
Trade-Offs among Adaptive Substitutions.” Evolution. Oxford University
Press, 2025. https://doi.org/10.1093/evolut/qpaf061.
ieee: T. Connallon et al., “Predicting the prevalence of genetic trade-offs
among adaptive substitutions,” Evolution, vol. 79, no. 7. Oxford University
Press, pp. 1243–1255, 2025.
ista: Connallon T, Czuppon P, Olito C, Goedert D, Kokko H, Nava-Bolaños A, Nilén
S, Svensson EI, Zwoinska M, Dutoit L, Ruzicka F. 2025. Predicting the prevalence
of genetic trade-offs among adaptive substitutions. Evolution. 79(7), 1243–1255.
mla: Connallon, Tim, et al. “Predicting the Prevalence of Genetic Trade-Offs among
Adaptive Substitutions.” Evolution, vol. 79, no. 7, Oxford University Press,
2025, pp. 1243–55, doi:10.1093/evolut/qpaf061.
short: T. Connallon, P. Czuppon, C. Olito, D. Goedert, H. Kokko, A. Nava-Bolaños,
S. Nilén, E.I. Svensson, M. Zwoinska, L. Dutoit, F. Ruzicka, Evolution 79 (2025)
1243–1255.
date_created: 2025-07-21T07:57:28Z
date_published: 2025-07-01T00:00:00Z
date_updated: 2025-09-30T14:06:38Z
day: '01'
ddc:
- '570'
department:
- _id: BeVi
doi: 10.1093/evolut/qpaf061
ec_funded: 1
external_id:
isi:
- '001477180800001'
file:
- access_level: open_access
checksum: 68c4c996d0e8c9ee3d4fb61bca75b31a
content_type: application/pdf
creator: dernst
date_created: 2025-07-22T10:04:57Z
date_updated: 2025-07-22T10:04:57Z
file_id: '20068'
file_name: 2025_Evolution_Connallon.pdf
file_size: 8150623
relation: main_file
success: 1
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has_accepted_license: '1'
intvolume: ' 79'
isi: 1
issue: '7'
language:
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month: '07'
oa: 1
oa_version: Published Version
page: 1243-1255
project:
- _id: fc2ed2f7-9c52-11eb-aca3-c01059dda49c
call_identifier: H2020
grant_number: '101034413'
name: 'IST-BRIDGE: International postdoctoral program'
publication: Evolution
publication_identifier:
eissn:
- 1558-5646
issn:
- 0014-3820
publication_status: published
publisher: Oxford University Press
quality_controlled: '1'
status: public
title: Predicting the prevalence of genetic trade-offs among adaptive substitutions
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: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 79
year: '2025'
...
---
OA_place: publisher
OA_type: hybrid
PlanS_conform: '1'
_id: '20655'
abstract:
- lang: eng
text: 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.
acknowledgement: 'We thank Brian Charlesworth, Deborah Charlesworth, and Sally Otto
for extensive comments and suggestions. We also thank Göran Arnqvist, Adam Eyre-Walker,
Philip Hedrick, Jitka Polechová, and Henrique Teotónio for further helpful comments
on the manuscript. This work was supported by a H2020 Marie Skłodowska-Curie COFUND
Action fellowship (#101034413, to F. R.), the Birgitta Sintring Foundation (#S2024-0007,
to M. K. Z.), the Research Council of Norway (302619, to D. G.), the Alexander von
Humboldt Foundation (to H. K.), the Swiss National Science Foundation (#211549,
to X. L. R.), the Swedish Research Council (#2022-03603, to CO; #2020-03123, to
E. I. S.) and the European Research Council (ERC-2023-STG-#101117517, to C. O.).
We are particularly grateful to the European Society for Evolutionary Biology for
funding a Special Topics Network workshop (to T. C., H. K., E. I. S.), from which
this review began. Open Access funding provided by Institute of Science and Technology
Austria/KEMÖ.'
article_number: brv.70103
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Filip
full_name: Ruzicka, Filip
id: 347955dd-57b0-11ee-9095-c28bdd368f4b
last_name: Ruzicka
- first_name: Martyna K.
full_name: Zwoinska, Martyna K.
last_name: Zwoinska
- first_name: Debora
full_name: Goedert, Debora
last_name: Goedert
- first_name: Hanna
full_name: Kokko, Hanna
last_name: Kokko
- first_name: Xiang‐Yi
full_name: Li Richter, Xiang‐Yi
last_name: Li Richter
- first_name: Iain R.
full_name: Moodie, Iain R.
last_name: Moodie
- first_name: Sofie
full_name: Nilén, Sofie
last_name: Nilén
- first_name: Colin
full_name: Olito, Colin
last_name: Olito
- first_name: Erik I.
full_name: Svensson, Erik I.
last_name: Svensson
- first_name: Peter
full_name: Czuppon, Peter
last_name: Czuppon
- first_name: Tim
full_name: Connallon, Tim
last_name: Connallon
citation:
ama: Ruzicka F, Zwoinska MK, Goedert D, et al. A century of theories of balancing
selection. Biological Reviews. 2025. doi:10.1111/brv.70103
apa: Ruzicka, F., Zwoinska, M. K., Goedert, D., Kokko, H., Li Richter, X., Moodie,
I. R., … Connallon, T. (2025). A century of theories of balancing selection. Biological
Reviews. Wiley. https://doi.org/10.1111/brv.70103
chicago: Ruzicka, Filip, Martyna K. Zwoinska, Debora Goedert, Hanna Kokko, Xiang‐Yi
Li Richter, Iain R. Moodie, Sofie Nilén, et al. “A Century of Theories of Balancing
Selection.” Biological Reviews. Wiley, 2025. https://doi.org/10.1111/brv.70103.
ieee: F. Ruzicka et al., “A century of theories of balancing selection,”
Biological Reviews. Wiley, 2025.
ista: Ruzicka F, Zwoinska MK, Goedert D, Kokko H, Li Richter X, Moodie IR, Nilén
S, Olito C, Svensson EI, Czuppon P, Connallon T. 2025. A century of theories of
balancing selection. Biological Reviews., brv. 70103.
mla: Ruzicka, Filip, et al. “A Century of Theories of Balancing Selection.” Biological
Reviews, brv. 70103, Wiley, 2025, doi:10.1111/brv.70103.
short: F. Ruzicka, M.K. Zwoinska, D. Goedert, H. Kokko, X. Li Richter, I.R. Moodie,
S. Nilén, C. Olito, E.I. Svensson, P. Czuppon, T. Connallon, Biological Reviews
(2025).
corr_author: '1'
date_created: 2025-11-19T09:43:50Z
date_published: 2025-11-14T00:00:00Z
date_updated: 2025-12-01T15:28:22Z
day: '14'
ddc:
- '570'
department:
- _id: BeVi
doi: 10.1111/brv.70103
ec_funded: 1
external_id:
isi:
- '001614285900001'
pmid:
- '41235821 '
has_accepted_license: '1'
isi: 1
language:
- iso: eng
main_file_link:
- open_access: '1'
url: https://doi.org/10.1111/brv.70103
month: '11'
oa: 1
oa_version: Published Version
pmid: 1
project:
- _id: fc2ed2f7-9c52-11eb-aca3-c01059dda49c
call_identifier: H2020
grant_number: '101034413'
name: 'IST-BRIDGE: International postdoctoral program'
publication: Biological Reviews
publication_identifier:
eissn:
- 1469-185X
issn:
- 1464-7931
publication_status: epub_ahead
publisher: Wiley
quality_controlled: '1'
scopus_import: '1'
status: public
title: A century of theories of balancing selection
tmp:
image: /images/cc_by_nc.png
legal_code_url: https://creativecommons.org/licenses/by-nc/4.0/legalcode
name: Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
short: CC BY-NC (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
year: '2025'
...
---
OA_place: publisher
OA_type: hybrid
_id: '18479'
abstract:
- lang: eng
text: The dominance of beneficial mutations is a key evolutionary parameter affecting
the rate and genetic basis of adaptation, yet it is notoriously difficult to estimate.
A leading method to infer it is to compare the relative rates of adaptive substitution
for X-linked and autosomal genes, which—according to a classic model by Charlesworth
et al. (1987)—is a simple function of the dominance of new beneficial mutations.
Recent evidence that rates of adaptive substitution are faster for X-linked genes
implies, accordingly, that beneficial mutations are usually recessive. However,
this conclusion is incompatible with leading theories of dominance, which predict
that beneficial mutations tend to be dominant or overdominant with respect to
fitness. To address this incompatibility, we use Fisher’s geometric model to predict
the distribution of fitness effects of new mutations and the relative rates of
positively selected substitution on the X and autosomes. Previous predictions
of faster-X theory emerge as a special case of our model in which the phenotypic
effects of mutations are small relative to the distance to the phenotypic optimum.
But as mutational effects become large relative to the optimum, we observe an
elevated tempo of positively selected substitutions on the X relative to the autosomes
across a broader range of dominance conditions, including those predicted by theories
of dominance. Our results imply that, contrary to previous models, dominant and
overdominant beneficial mutations can plausibly generate patterns of faster-X
adaptation. We discuss resulting implications for genomic studies of adaptation
and inferences of dominance.
acknowledgement: This work was supported by funds from the Australian Research Council
and The School of Biological Sciences at Monash University. F.R. was funded by a
H2020 Marie Skłodowska-Curie COFUND Action (No. 101034413). We thank three anonymous
reviewers for suggestions that substantially improved the paper and breadth of the
analysis.
article_number: e2406335121
article_processing_charge: Yes (in subscription journal)
article_type: original
author:
- first_name: Yasmine
full_name: Mcdonough, Yasmine
last_name: Mcdonough
- first_name: Filip
full_name: Ruzicka, Filip
id: 347955dd-57b0-11ee-9095-c28bdd368f4b
last_name: Ruzicka
- first_name: Tim
full_name: Connallon, Tim
last_name: Connallon
citation:
ama: Mcdonough Y, Ruzicka F, Connallon T. Reconciling theories of dominance with
the relative rates of adaptive substitution on sex chromosomes and autosomes.
Proceedings of the National Academy of Sciences of the United States of America.
2024;121(44). doi:10.1073/pnas.2406335121
apa: Mcdonough, Y., Ruzicka, F., & Connallon, T. (2024). Reconciling theories
of dominance with the relative rates of adaptive substitution on sex chromosomes
and autosomes. Proceedings of the National Academy of Sciences of the United
States of America. National Academy of Sciences. https://doi.org/10.1073/pnas.2406335121
chicago: Mcdonough, Yasmine, Filip Ruzicka, and Tim Connallon. “Reconciling Theories
of Dominance with the Relative Rates of Adaptive Substitution on Sex Chromosomes
and Autosomes.” Proceedings of the National Academy of Sciences of the United
States of America. National Academy of Sciences, 2024. https://doi.org/10.1073/pnas.2406335121.
ieee: Y. Mcdonough, F. Ruzicka, and T. Connallon, “Reconciling theories of dominance
with the relative rates of adaptive substitution on sex chromosomes and autosomes,”
Proceedings of the National Academy of Sciences of the United States of America,
vol. 121, no. 44. National Academy of Sciences, 2024.
ista: Mcdonough Y, Ruzicka F, Connallon T. 2024. Reconciling theories of dominance
with the relative rates of adaptive substitution on sex chromosomes and autosomes.
Proceedings of the National Academy of Sciences of the United States of America.
121(44), e2406335121.
mla: Mcdonough, Yasmine, et al. “Reconciling Theories of Dominance with the Relative
Rates of Adaptive Substitution on Sex Chromosomes and Autosomes.” Proceedings
of the National Academy of Sciences of the United States of America, vol.
121, no. 44, e2406335121, National Academy of Sciences, 2024, doi:10.1073/pnas.2406335121.
short: Y. Mcdonough, F. Ruzicka, T. Connallon, Proceedings of the National Academy
of Sciences of the United States of America 121 (2024).
date_created: 2024-10-27T23:01:44Z
date_published: 2024-10-29T00:00:00Z
date_updated: 2025-09-08T14:31:58Z
day: '29'
ddc:
- '570'
department:
- _id: BeVi
doi: 10.1073/pnas.2406335121
ec_funded: 1
external_id:
isi:
- '001359216400017'
pmid:
- '39436652'
file:
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checksum: 73db3c87b35753e0f4324417f164a35e
content_type: application/pdf
creator: dernst
date_created: 2024-11-04T10:29:43Z
date_updated: 2024-11-04T10:29:43Z
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file_name: 2024_PNAS_McDonough.pdf
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relation: main_file
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file_date_updated: 2024-11-04T10:29:43Z
has_accepted_license: '1'
intvolume: ' 121'
isi: 1
issue: '44'
language:
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month: '10'
oa: 1
oa_version: Published Version
pmid: 1
project:
- _id: fc2ed2f7-9c52-11eb-aca3-c01059dda49c
call_identifier: H2020
grant_number: '101034413'
name: 'IST-BRIDGE: International postdoctoral program'
publication: Proceedings of the National Academy of Sciences of the United States
of America
publication_identifier:
eissn:
- 1091-6490
publication_status: published
publisher: National Academy of Sciences
quality_controlled: '1'
scopus_import: '1'
status: public
title: Reconciling theories of dominance with the relative rates of adaptive substitution
on sex chromosomes and autosomes
tmp:
image: /images/cc_by_nc_nd.png
legal_code_url: https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode
name: Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International
(CC BY-NC-ND 4.0)
short: CC BY-NC-ND (4.0)
type: journal_article
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 121
year: '2024'
...