The DOI matches its Crossref record; 61 of the 61 references it cites were found in a registry or an index.
Mathematical Elegance with Biochemical Realism: The Covarion Model of Molecular Evolution
Open accessAbstract
There is an apparent paradox in our understanding of molecular evolution. Current biochemically based models predict that evolutionary trees should not be recoverable for divergences beyond a few hundred million years. In practice, however, trees often appear to be recovered from much older times. Mathematical models, such as those assuming that sites evolve at different rates [including a Gamma distribution of rates across sites (RAS)] may in theory allow the recovery of some ancient divergences. However, such models require that each site maintain its characteristic rate over the whole evolutionary period. This assumption, however, contradicts the knowledge that tertiary structures diverge with time, invalidating the rate-constancy assumption of purely mathematical models. We report here that a hidden Markov version of the covarion model can meet both biochemical and statistical requirements for the analysis of sequence data. The model was proposed on biochemical grounds and can be implemented with only two additional parameters. The two hidden parts of this model are the proportion of sites free to vary (covarions) and the rate of interchange between fixed sites and these variable sites. Simulation results are consistent with this approach, providing a better framework for understanding anciently diverged sequences than the standard RAS models. However, a Gamma distribution of rates may approximate a covarion model and may possibly be justified on these grounds. The accurate reconstruction of older divergences from sequence data is still a major problem, and molecular evolution still requires mathematical models that also have a sound biochemical basis.
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“According to this model, the rate of gene evolution is governed by the overall mutation rate and the proportion of sites at which changes are neutral 6 (FIG. 1b).”
From The modern molecular clock (2003) · cites this paper as [6]
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Simon, C. M., Frati, F., Beckenbach, A. T., Crespi, B., Liu, H., & Flook, P. K. (1994) · Annals of the Entomological Society of America · doi:10.1093/aesa/87.6.651 · Cited by 6,119
Conjectures and Refutations: The Growth of Scientific Knowledge.
Hutchison, T. W., & Popper, K. R. (1964) · Economica · doi:10.2307/2551371 · Cited by 4,232
Conjectures and Refutations: The Growth of Scientific Knowledge.
HESSE, M. B., & Popper, K. R. (1965) · The Philosophical Quarterly · doi:10.2307/2218271 · Cited by 4,213
The relation between the divergence of sequence and structure in proteins.
Chothia, C., & Lesk, A. M. (1986) · The EMBO Journal · doi:10.1002/j.1460-2075.1986.tb04288.x · Cited by 2,458
A codon-based model of nucleotide substitution for protein-coding DNA sequences.
(1994) · Molecular Biology and Evolution · doi:10.1093/oxfordjournals.molbev.a040153 · Cited by 1,960
Hidden Markov Models in Computational Biology
Krogh, A., Brown, M., Mian, S. I., Sjölander, K. V., & Haussler, D. H. (1994) · Journal of Molecular Biology · doi:10.1006/jmbi.1994.1104 · Cited by 1,948
Evolutionarily Conserved Pathways of Energetic Connectivity in Protein Families
Lockless, S. W., & Ranganathan, R. (1999) · Science · doi:10.1126/science.286.5438.295 · Cited by 1,288
Among-site rate variation and its impact on phylogenetic analyses
Yang, Z. (1996) · Trends in Ecology & Evolution · doi:10.1016/0169-5347(96)10041-0 · Cited by 1,202
Molecular Evolution: A Phylogenetic Approach
Page, R., & Holmes, E. C. (1998) · Medical Entomology and Zoology · Cited by 984
A Hidden Markov Model approach to variation among sites in rate of evolution
Felsenstein, J., & Churchill, G. A. (1996) · Molecular Biology and Evolution · doi:10.1093/oxfordjournals.molbev.a025575 · Cited by 846
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Metadata
- Authors
- Penny, D., McComish, B. J., Charleston, M. A., & Hendy, M. D.
- Affiliations
- Massey University (NZ)
- Venue
- Journal of Molecular Evolution
- Published
- 2001-12
- Volume
- 53
- Issue
- 6
- Pages
- 711-723
- ISSN
- 0022-2844 (ISSN-L)
- 1432-1432
- OpenAlex
- W2139677507
- Cited by
- 287
- References
- 61
- Citations by year
- 2026: 7 · 2025: 44 · 2024: 63 · 2023: 5 · 2022: 4 · 2021: 5 · 2020: 6 · 2019: 6 · 2018: 6 · 2017: 4 · 2016: 4 · 2015: 4 · 2014: 11 · 2013: 5 · 2012: 11
- FWCI
- 3.70
- Type
- journal article
- Language
- en
- Open access
- green
- figshare.com
- Licence
- Locations
- Journal of Molecular Evolution (publishedVersion)
- PubMed (publishedVersion)
- eCite Digital Repository (University of Tasmania) (submittedVersion)
- Figshare (submittedVersion, cc-by)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Phylogenetic Studies
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Evolution and Genetic Dynamics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › RNA and protein synthesis mechanisms
- Keywords
- hidden Markov models, molecular evolution, phylogenetic inference
- MeSH terms
- Models, Biological, Models, Chemical, Evolution, Molecular
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