---
OA_place: publisher
OA_type: hybrid
_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: 804-825
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. 2026;101(2). doi:10.1111/brv.70103
apa: Ruzicka, F., Zwoinska, M. K., Goedert, D., Kokko, H., Li Richter, X., Moodie,
I. R., … Connallon, T. (2026). 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, 2026. https://doi.org/10.1111/brv.70103.
ieee: F. Ruzicka et al., “A century of theories of balancing selection,”
Biological Reviews, vol. 101, no. 2. Wiley, 2026.
ista: Ruzicka F, Zwoinska MK, Goedert D, Kokko H, Li Richter X, Moodie IR, Nilén
S, Olito C, Svensson EI, Czuppon P, Connallon T. 2026. A century of theories of
balancing selection. Biological Reviews. 101(2), 804–825.
mla: Ruzicka, Filip, et al. “A Century of Theories of Balancing Selection.” Biological
Reviews, vol. 101, no. 2, 804–825, Wiley, 2026, 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
101 (2026).
corr_author: '1'
das_tickbox: '0'
date_created: 2025-11-19T09:43:50Z
date_published: 2026-04-01T00:00:00Z
date_updated: 2026-07-27T08:08:09Z
day: '01'
ddc:
- '570'
department:
- _id: BeVi
doi: 10.1111/brv.70103
ec_funded: 1
external_id:
isi:
- '001614285900001'
pmid:
- '41235821 '
file:
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checksum: 167d95cf0570d6e3653ab349b2a4355c
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success: 1
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fulldoi: https://doi.org/10.1111/brv.70103
has_accepted_license: '1'
intvolume: ' 101'
isi: 1
issue: '2'
keyword:
- evolutionary theory
- population genetics
- balancing selection
- heterozygote advantage
- trade-offs
- negative frequency-dependent selection
- fitness variation
- mathematical modelling
language:
- iso: eng
license: https://creativecommons.org/licenses/by-nc/4.0/
month: '04'
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: published
publisher: Wiley
quality_controlled: '1'
researchdata_availability: no
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supplementarymaterial: yes
title: A century of theories of balancing selection
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...
---
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abstract:
- lang: eng
text: "Sex chromosomes and autosomes exhibit very different evolutionary dynamics.\r\nThe
Y chromosome usually degenerates, leaving many X-linked loci hemizygous in\r\nmales.
Since recessive X-linked mutations are always exposed to selection in males,\r\nselection
is more efficient on the X chromosome than on autosomes on recessive\r\nmutations,
leading to faster adaptation on the X chromosome than other genomic\r\nregions,
if beneficial mutations are on average recessive (known as the Faster-X\r\neffect).
In the presence of the functional, but non-recombining gametolog on the Y (as\r\nis
often the case in young non-recombining regions), recessive mutations are\r\nsheltered
from selection on the X chromosome. We model this scenario and show that\r\nthe
efficiency of selection is reduced on diploid X loci due to sheltering by the
Y\r\nchromosome. Reduced efficiency of selection leads to slower adaptation and\r\nincreased
accumulation of deleterious mutations (Slower-X effect). We extended this\r\nmodel
to explore the effect of sex-specific selection on degeneration of sex\r\nchromosomes,
showing theoretically that male-limited genes degenerate on the X\r\nchromosome
and female-biased genes degenerate on the Y chromosome. This\r\nprediction depends
on the effective population size and the mutation rate, explaining\r\nthe variety
of sex chromosome degeneration patterns observed in nature.\r\nTo test for direct
evidence of a Slower-X (or Slower-Z) effect, we analyzed the\r\nZW sex chromosomes
of the flatworm Schistosoma japonicum, which have a very\r\nyoung non-recombining
region with non-degenerated W. Diploid Z-linked genes have\r\nhigher ratios of
non-synonymous to synonymous polymorphisms than autosomal\r\ngenes, supporting
reduced efficiency of selection on the diploid Z region. These results\r\nprovide
evidence of sheltering by the W chromosome, a mechanism that could\r\ncontribute
to Z (X) chromosome degeneration, and illustrate contrasting evolutionary\r\npatterns
in old and young sex chromosome regions. In addition, genes with sexspecific patterns
of expression show opposite patterns of selection in the young\r\n(diploid) and
old (hemizygous) Z, showing the complex manner in which sex-specific selection
shapes the evolutionary patterns of sex chromosomes. "
acknowledged_ssus:
- _id: ScienComp
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alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Andrea
full_name: Mrnjavac, Andrea
id: 353FAC84-AE61-11E9-8BFC-00D3E5697425
last_name: Mrnjavac
citation:
ama: Mrnjavac A. Early stages of sex chromosome evolution. 2024. doi:10.15479/at:ista:18531
apa: Mrnjavac, A. (2024). Early stages of sex chromosome evolution. Institute
of Science and Technology Austria. https://doi.org/10.15479/at:ista:18531
chicago: Mrnjavac, Andrea. “Early Stages of Sex Chromosome Evolution.” Institute
of Science and Technology Austria, 2024. https://doi.org/10.15479/at:ista:18531.
ieee: A. Mrnjavac, “Early stages of sex chromosome evolution,” Institute of Science
and Technology Austria, 2024.
ista: Mrnjavac A. 2024. Early stages of sex chromosome evolution. Institute of Science
and Technology Austria.
mla: Mrnjavac, Andrea. Early Stages of Sex Chromosome Evolution. Institute
of Science and Technology Austria, 2024, doi:10.15479/at:ista:18531.
short: A. Mrnjavac, Early Stages of Sex Chromosome Evolution, Institute of Science
and Technology Austria, 2024.
corr_author: '1'
date_created: 2024-11-11T08:40:45Z
date_published: 2024-11-11T00:00:00Z
date_updated: 2026-10-02T11:14:08Z
day: '11'
ddc:
- '576'
degree_awarded: PhD
department:
- _id: GradSch
- _id: BeVi
doi: 10.15479/at:ista:18531
doi_confirm: '1'
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file_date_updated: 2025-05-11T22:30:04Z
fulldoi: https://doi.org/10.15479/at:ista:18531
has_accepted_license: '1'
keyword:
- Sex chromosomes
- evolution
- selection
- sheltering
language:
- iso: eng
license: https://creativecommons.org/licenses/by-nc-nd/4.0/
month: '11'
oa: 1
oa_version: Published Version
page: '181'
publication_identifier:
issn:
- 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
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researchdata_availability: unclear
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supervisor:
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
supplementarymaterial: unclear
title: Early stages of sex chromosome evolution
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short: CC BY-NC-ND (4.0)
type: dissertation
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
year: '2024'
...
---
_id: '5559'
abstract:
- lang: eng
text: Strong amplifiers of natural selection
article_processing_charge: No
author:
- first_name: Andreas
full_name: Pavlogiannis, Andreas
id: 49704004-F248-11E8-B48F-1D18A9856A87
last_name: Pavlogiannis
orcid: 0000-0002-8943-0722
- first_name: Josef
full_name: Tkadlec, Josef
id: 3F24CCC8-F248-11E8-B48F-1D18A9856A87
last_name: Tkadlec
orcid: 0000-0002-1097-9684
- first_name: Krishnendu
full_name: Chatterjee, Krishnendu
id: 2E5DCA20-F248-11E8-B48F-1D18A9856A87
last_name: Chatterjee
orcid: 0000-0002-4561-241X
- first_name: Martin
full_name: Nowak , Martin
last_name: 'Nowak '
citation:
ama: Pavlogiannis A, Tkadlec J, Chatterjee K, Nowak M. Strong amplifiers of natural
selection. 2017. doi:10.15479/AT:ISTA:51
apa: Pavlogiannis, A., Tkadlec, J., Chatterjee, K., & Nowak , M. (2017). Strong
amplifiers of natural selection. Institute of Science and Technology Austria.
https://doi.org/10.15479/AT:ISTA:51
chicago: Pavlogiannis, Andreas, Josef Tkadlec, Krishnendu Chatterjee, and Martin
Nowak . “Strong Amplifiers of Natural Selection.” Institute of Science and Technology
Austria, 2017. https://doi.org/10.15479/AT:ISTA:51.
ieee: A. Pavlogiannis, J. Tkadlec, K. Chatterjee, and M. Nowak , “Strong amplifiers
of natural selection.” Institute of Science and Technology Austria, 2017.
ista: Pavlogiannis A, Tkadlec J, Chatterjee K, Nowak M. 2017. Strong amplifiers
of natural selection, Institute of Science and Technology Austria, 10.15479/AT:ISTA:51.
mla: Pavlogiannis, Andreas, et al. Strong Amplifiers of Natural Selection.
Institute of Science and Technology Austria, 2017, doi:10.15479/AT:ISTA:51.
short: A. Pavlogiannis, J. Tkadlec, K. Chatterjee, M. Nowak , (2017).
datarep_id: '51'
date_created: 2018-12-12T12:31:32Z
date_published: 2017-01-02T00:00:00Z
date_updated: 2025-04-15T08:12:19Z
day: '02'
ddc:
- '519'
department:
- _id: KrCh
doi: 10.15479/AT:ISTA:51
ec_funded: 1
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fulldoi: https://doi.org/10.15479/AT:ISTA:51
has_accepted_license: '1'
keyword:
- natural selection
month: '01'
oa: 1
oa_version: Published Version
project:
- _id: 2581B60A-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '279307'
name: 'Quantitative Graph Games: Theory and Applications'
publisher: Institute of Science and Technology Austria
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relation: research_paper
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title: Strong amplifiers of natural selection
type: research_data
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year: '2017'
...
---
_id: '5553'
abstract:
- lang: eng
text: "Genotypic, phenotypic and demographic data for 2128 wild snapdragons and
1127 open-pollinated progeny from a natural hybrid zone, collected as part of
Tom Ellis' PhD thesis (submitted) February 2016).\r\n\r\nTissue samples were sent
to LGC Genomics in Berlin for DNA extraction, and genotyping at 70 SNP markers
by KASPR genotyping. 29 of these SNPs failed to amplify reliably, and have been
removed from this dataset.\r\n\r\nOther data were retreived from an online database
of this population at www.antspec.org."
article_processing_charge: No
author:
- first_name: David
full_name: Field, David
id: 419049E2-F248-11E8-B48F-1D18A9856A87
last_name: Field
orcid: 0000-0002-4014-8478
- first_name: Thomas
full_name: Ellis, Thomas
id: 3153D6D4-F248-11E8-B48F-1D18A9856A87
last_name: Ellis
orcid: 0000-0002-8511-0254
citation:
ama: Field D, Ellis T. Inference of mating patterns among wild snapdragons in a
natural hybrid zone in 2012. 2016. doi:10.15479/AT:ISTA:37
apa: Field, D., & Ellis, T. (2016). Inference of mating patterns among wild
snapdragons in a natural hybrid zone in 2012. Institute of Science and Technology
Austria. https://doi.org/10.15479/AT:ISTA:37
chicago: Field, David, and Thomas Ellis. “Inference of Mating Patterns among Wild
Snapdragons in a Natural Hybrid Zone in 2012.” Institute of Science and Technology
Austria, 2016. https://doi.org/10.15479/AT:ISTA:37.
ieee: D. Field and T. Ellis, “Inference of mating patterns among wild snapdragons
in a natural hybrid zone in 2012.” Institute of Science and Technology Austria,
2016.
ista: Field D, Ellis T. 2016. Inference of mating patterns among wild snapdragons
in a natural hybrid zone in 2012, Institute of Science and Technology Austria,
10.15479/AT:ISTA:37.
mla: Field, David, and Thomas Ellis. Inference of Mating Patterns among Wild
Snapdragons in a Natural Hybrid Zone in 2012. Institute of Science and Technology
Austria, 2016, doi:10.15479/AT:ISTA:37.
short: D. Field, T. Ellis, (2016).
contributor:
- contributor_type: project_manager
first_name: Nicholas H
id: 4880FE40-F248-11E8-B48F-1D18A9856A87
last_name: Barton
orcid: 0000-0002-8548-5240
datarep_id: '37'
date_created: 2018-12-12T12:31:30Z
date_published: 2016-02-19T00:00:00Z
date_updated: 2026-07-30T14:56:13Z
day: '19'
ddc:
- '576'
department:
- _id: NiBa
doi: 10.15479/AT:ISTA:37
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creator: system
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fulldoi: https://doi.org/10.15479/AT:ISTA:37
has_accepted_license: '1'
keyword:
- paternity assignment
- pedigree
- matting patterns
- assortative mating
- Antirrhinum majus
- frequency-dependent selection
- plant-pollinator interaction
license: https://creativecommons.org/publicdomain/zero/1.0/
month: '02'
oa: 1
oa_version: Published Version
publisher: Institute of Science and Technology Austria
related_material:
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status: public
title: Inference of mating patterns among wild snapdragons in a natural hybrid zone
in 2012
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---
_id: '10899'
article_processing_charge: No
author:
- first_name: Nicholas H
full_name: Barton, Nicholas H
id: 4880FE40-F248-11E8-B48F-1D18A9856A87
last_name: Barton
orcid: 0000-0002-8548-5240
citation:
ama: 'Barton NH. Differentiation. In: Encyclopedia of Biodiversity. 2nd ed.
Elsevier; 2013:508-515. doi:10.1016/b978-0-12-384719-5.00031-9'
apa: Barton, N. H. (2013). Differentiation. In Encyclopedia of Biodiversity
(2nd ed., pp. 508–515). Elsevier. https://doi.org/10.1016/b978-0-12-384719-5.00031-9
chicago: Barton, Nicholas H. “Differentiation.” In Encyclopedia of Biodiversity,
2nd ed., 508–15. Elsevier, 2013. https://doi.org/10.1016/b978-0-12-384719-5.00031-9.
ieee: N. H. Barton, “Differentiation,” in Encyclopedia of Biodiversity, 2nd
ed., Elsevier, 2013, pp. 508–515.
ista: 'Barton NH. 2013.Differentiation. In: Encyclopedia of Biodiversity. , 508–515.'
mla: Barton, Nicholas H. “Differentiation.” Encyclopedia of Biodiversity,
2nd ed., Elsevier, 2013, pp. 508–15, doi:10.1016/b978-0-12-384719-5.00031-9.
short: N.H. Barton, in:, Encyclopedia of Biodiversity, 2nd ed., Elsevier, 2013,
pp. 508–515.
corr_author: '1'
date_created: 2022-03-21T07:46:22Z
date_published: 2013-01-01T00:00:00Z
date_updated: 2024-10-09T21:02:37Z
day: '01'
department:
- _id: NiBa
doi: 10.1016/b978-0-12-384719-5.00031-9
edition: '2'
fulldoi: https://doi.org/10.1016/b978-0-12-384719-5.00031-9
keyword:
- Adaptive landscape
- Cline
- Coalescent process
- Gene flow
- Hybrid zone
- Local adaptation
- Natural selection
- Neutral theory
- Population structure
- Speciation
language:
- iso: eng
month: '01'
oa_version: None
page: 508-515
publication: Encyclopedia of Biodiversity
publication_identifier:
isbn:
- 978-0-12-384720-1
publication_status: published
publisher: Elsevier
quality_controlled: '1'
scopus_import: '1'
status: public
title: Differentiation
type: book_chapter
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
year: '2013'
...
---
OA_type: closed access
_id: '869'
abstract:
- lang: eng
text: The impact of synonymous nucleotide substitutions on fitness in mammals remains
controversial. Despite some indications of selective constraint, synonymous sites
are often assumed to be neutral, and the rate of their evolution is used as a
proxy for mutation rate. We subdivide all sites into four classes in terms of
the mutable CpG context, nonCpG, postC, preG, and postCpreG, and compare four-fold
synonymous sites and intron sites residing outside transposable elements. The
distribution of the rate of evolution across all synonymous sites is trimodal.
Rate of evolution at nonCpG synonymous sites, not preceded by C and not followed
by G, is ∼10% below that at such intron sites. In contrast, rate of evolution
at postCpreG synonymous sites is ∼30% above that at such intron sites. Finally,
synonymous and intron postC and preG sites evolve at similar rates. The relationship
between the levels of polymorphism at the corresponding synonymous and intron
sites is very similar to that between their rates of evolution. Within every class,
synonymous sites are occupied by G or C much more often than intron sites, whose
nucleotide composition is consistent with neutral mutation-drift equilibrium.
These patterns suggest that synonymous sites are under weak selection in favor
of G and C, with the average coefficient s∼0.25/Ne∼10-5, where Ne is the effective
population size. Such selection decelerates evolution and reduces variability
at sites with symmetric mutation, but has the opposite effects at sites where
the favored nucleotides are more mutable. The amino-acid composition of proteins
dictates that many synonymous sites are CpGprone, which causes them, on average,
to evolve faster and to be more polymorphic than intron sites. An average genotype
carries ∼107 suboptimal nucleotides at synonymous sites, implying synergistic
epistasis in selection against them.
acknowledgement: This research was supported in part by the Intramural Research Program
of the NIH, National Library of Medicine.
article_processing_charge: No
article_type: original
author:
- first_name: Fyodor
full_name: Kondrashov, Fyodor
id: 44FDEF62-F248-11E8-B48F-1D18A9856A87
last_name: Kondrashov
orcid: 0000-0001-8243-4694
- first_name: Aleksey
full_name: Ogurtsov, Aleksey
last_name: Ogurtsov
- first_name: Alexey
full_name: Kondrashov, Alexey
last_name: Kondrashov
citation:
ama: Kondrashov F, Ogurtsov A, Kondrashov A. Selection in favor of nucleotides G
and C diversifies evolution rates and levels of polymorphism at mammalian synonymous
sites. Journal of Theoretical Biology. 2006;240(4):616-626. doi:10.1016/j.jtbi.2005.10.020
apa: Kondrashov, F., Ogurtsov, A., & Kondrashov, A. (2006). Selection in favor
of nucleotides G and C diversifies evolution rates and levels of polymorphism
at mammalian synonymous sites. Journal of Theoretical Biology. Elsevier.
https://doi.org/10.1016/j.jtbi.2005.10.020
chicago: Kondrashov, Fyodor, Aleksey Ogurtsov, and Alexey Kondrashov. “Selection
in Favor of Nucleotides G and C Diversifies Evolution Rates and Levels of Polymorphism
at Mammalian Synonymous Sites.” Journal of Theoretical Biology. Elsevier,
2006. https://doi.org/10.1016/j.jtbi.2005.10.020.
ieee: F. Kondrashov, A. Ogurtsov, and A. Kondrashov, “Selection in favor of nucleotides
G and C diversifies evolution rates and levels of polymorphism at mammalian synonymous
sites,” Journal of Theoretical Biology, vol. 240, no. 4. Elsevier, pp.
616–626, 2006.
ista: Kondrashov F, Ogurtsov A, Kondrashov A. 2006. Selection in favor of nucleotides
G and C diversifies evolution rates and levels of polymorphism at mammalian synonymous
sites. Journal of Theoretical Biology. 240(4), 616–626.
mla: Kondrashov, Fyodor, et al. “Selection in Favor of Nucleotides G and C Diversifies
Evolution Rates and Levels of Polymorphism at Mammalian Synonymous Sites.” Journal
of Theoretical Biology, vol. 240, no. 4, Elsevier, 2006, pp. 616–26, doi:10.1016/j.jtbi.2005.10.020.
short: F. Kondrashov, A. Ogurtsov, A. Kondrashov, Journal of Theoretical Biology
240 (2006) 616–626.
date_created: 2018-12-11T11:48:56Z
date_published: 2006-06-21T00:00:00Z
date_updated: 2026-05-08T10:18:03Z
day: '21'
doi: 10.1016/j.jtbi.2005.10.020
extern: '1'
external_id:
pmid:
- '16343547'
fulldoi: https://doi.org/10.1016/j.jtbi.2005.10.020
intvolume: ' 240'
issue: '4'
keyword:
- Mutation
- Selection
- Synonymous site
- Evolution
- Genetic drift
language:
- iso: eng
month: '06'
oa_version: None
page: 616 - 626
pmid: 1
publication: Journal of Theoretical Biology
publication_identifier:
issn:
- 1095-8541
publication_status: published
publisher: Elsevier
publist_id: '6779'
quality_controlled: '1'
scopus_import: '1'
status: public
title: Selection in favor of nucleotides G and C diversifies evolution rates and levels
of polymorphism at mammalian synonymous sites
type: journal_article
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
volume: 240
year: '2006'
...