Origin and evolution of the genetic code: The universal enigma

Koonin, Eugene V.; Novozhilov, Artem S.

doi:10.1002/iub.146

Cited by 392RNA and protein synthesis mechanismsOrigins and Evolution of LifeEvolution and Genetic Dynamics

Origin and evolution of the genetic code: The universal enigma

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Koonin, Eugene V., Novozhilov, Artem S.·2009·IUBMB Life

DOI 10.1002/iub.146PMC PMC3293468

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Abstract

The genetic code is nearly universal, and the arrangement of the codons in the standard codon table is highly nonrandom. The three main concepts on the origin and evolution of the code are the stereochemical theory, according to which codon assignments are dictated by physicochemical affinity between amino acids and the cognate codons (anticodons); the coevolution theory, which posits that the code structure coevolved with amino acid biosynthesis pathways; and the error minimization theory under which selection to minimize the adverse effect of point mutations and translation errors was the principal factor of the code's evolution. These theories are not mutually exclusive and are also compatible with the frozen accident hypothesis, that is, the notion that the standard code might have no special properties but was fixed simply because all extant life forms share a common ancestor, with subsequent changes to the code, mostly, precluded by the deleterious effect of codon reassignment. Mathematical analysis of the structure and possible evolutionary trajectories of the code shows that it is highly robust to translational misreading but there are numerous more robust codes, so the standard code potentially could evolve from a random code via a short sequence of codon series reassignments. Thus, much of the evolution that led to the standard code could be a combination of frozen accident with selection for error minimization although contributions from coevolution of the code with metabolic pathways and weak affinities between amino acids and nucleotide triplets cannot be ruled out. However, such scenarios for the code evolution are based on formal schemes whose relevance to the actual primordial evolution is uncertain. A real understanding of the code origin and evolution is likely to be attainable only in conjunction with a credible scenario for the evolution of the coding principle itself and the translation system. © 2008 IUBMB IUBMB Life, 61(2): 99–111, 2009

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The principal debate revolves around four competing perspectives on the code's origin and evolution.
“The principal debate revolves around four competing perspectives on the code's origin and evolution (18, 39, 44).”
From Origin and Evolution of the Universal Genetic Code (2017) · cites this paper as [44]
The Introduction, under “The basic theories of the code nature, origin and evolution,” identifies three main theories and says they are not necessarily mutually exclusive. The paper also discusses the frozen-accident view as a default explanation, but does not present four competing perspectives as the principal debate. Page numbers were not available in our copy. (LLM, conf=0.88)
Checked by a reasoning model — may be wrong.
Experimental — accuracy not yet measured
We currently have no formal models for genetic code evolution.
“We currently have no such formal models for genetic code evolution [11,12,25,27].”
From On universal common ancestry, sequence similarity, and phylogenetic structure: the sins of P-values and the virtues of Bayesian evidence (2011) · cites this paper as [12]
The section “The adaptive theory: evidence of evolutionary optimization of the code” describes the Ardell–Sella model of code evolution. In “Instead of conclusions,” the authors say that studies of code evolution use formalized settings; their concern is whether those models reflect primordial evolution, not that formal models do not exist. Page numbers are unavailable in our copy. (LLM, conf=0.90)
Checked by a reasoning model — may be wrong.
Experimental — accuracy not yet measured
Given genes and proteins, the origin of molecular machines such as the ATPase is an impressive but not conceptually challenging evolutionary step, far less problematic than the hurdles at the origin of the ribosome and the code.
“Given genes and proteins, the origin of molecular machines such as the ATPase is an admittedly impressive, but not conceptually challenging, evolutionary step; it is a far less problematic increment than the hurdles that had to be surmounted at the origin of the ribosome and the code [110,117].”
From Early bioenergetic evolution (2013) · cites this paper as [110]
In “Instead of conclusions: How did the code evolve (and will we ever know)?” (page not available), the paper calls the origin of coding and the translation system a formidable problem and describes a circular difficulty: functional proteins require translation, while the evolution of translation needs a selective force. It does not discuss ATPase or say that, once genes and proteins exist, molecular machines are conceptually easier to evolve than the ribosome and code. (LLM, conf=0.84)
Experimental — accuracy not yet measured
The scenarios of code evolution do not have to be mutually exclusive.
“Clearly, these scenarios of code evolution do not have to be mutually exclusive, further complicating the problem (44).”
From Origin and Evolution of the Universal Genetic Code (2017) · cites this paper as [44]
The structure of the SGC is manifestly regular with respect to multiple criteria.
“The structure of the SGC-that is, the mapping of 64 codons to 20 amino acids and the stop signal-is manifestly regular with respect to multiple criteria (44, 84).”
From Origin and Evolution of the Universal Genetic Code (2017) · cites this paper as [44]

The standard genetic code is far from optimal because billions of possible code variants are even more robust to error.
“However, the SGC is far from being optimal because, given the enormous overall number of possible codes (>1084), billions of variants are even more robust to error [10].”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The three major concepts that strive to explain regularities in the code are the stereochemical, coevolution, and error minimization theories.
“The three major concepts that strive to explain the regularities in the code are the stereochemical, coevolution, and error minimization ‘theories’ [10,43,44].”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The stereochemical theory postulates that the structure of the code is determined by physicochemical affinity between amino acids and codons or anticodons.
“The stereochemical theory postulates that the structure of the code is determined by physicoo-chemical affinity between amino acids and codons or anticodons.”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The structure of the standard genetic code is clearly non-random by multiple criteria.
“The structure of the SGC, i.e., the mapping of 64 codons to 20 amino acids and the stop signal, is clearly non-random by multiple criteria [9,10].”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
Stereochemical affinity between amino acids and cognate codons or anticodons does not seem to account for the origin and evolution of the code.
“In particular, stereochemical affinity between amino acids and the cognate codons or anticodons does not seem to account for the origin and evolution of the code.”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The basic structure of the genetic code and the majority of codon assignments are truly universal.
“the basic structure of the code and the majority of the codon assignments are truly universal [10,11].”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
Quantitative analyses using cost functions have confirmed exceptional robustness of the standard genetic code: the probability to reach the same level of error minimization by random permutation is below 10^-6.
“Quantitative analyses of the code using cost functions derived from physico-chemical properties of amino acids or their evolutionary exchangeability have confirmed the exceptional robustness of the standard genetic code (SGC): the probability to reach the same level of error minimization as in the SGC by random permutation of codons is below 10−6 [10,13,14,15].”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The stereochemical evidence does not appear to be compelling; significant affinity for cognate triplets has been demonstrated only for five amino acids, and the triplet could be either codon or anticodon; all aptamer results compatible with affinity pertain to complex amino acids unlikely to have been available at code emergence.
“However, as discussed elsewhere [10,45], the stereochemical evidence does not appear to be compelling. The principal problems are two-fold. First, statistically significant affinity for cognate triplets has been demonstrated only for five amino acids, and the triplet could be either codon or anticodon [55]. Second, and more damning, all the results of aptamer experiments that appear compatible”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]
The error minimization theory holds that the structure of the code is determined by selection for robustness to errors.
“Hence the error minimization theory of code evolution, under which the structure of the code is determined by selection for robustness to errors.”
From Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code (2017) · cites this paper as [10]

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Evolution of the Genetic Triplet Code via Two Types of Doublet Codons
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The triplet genetic code had a doublet predecessor
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Question 6: Coevolution Theory of the Genetic Code: A Proven Theory
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Codon Usage Decreases the Error Minimization Within the Genetic Code
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Error Minimization and Coding Triplet/Binding Site Associations Are Independent Features of the Canonical Genetic Code
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A Comparative Genomics Analysis of Codon Reassignments Reveals a Link with Mitochondrial Proteome Size and a Mechanism of Genetic Code Change Via Suppressor tRNAs
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Block structure and stability of the genetic code
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Generic Darwinian selection in catalytic protocell assemblies
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A Blind Empiricism Against the Coevolution Theory of the Origin of the Genetic Code
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Universality in the Genetic Code
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The Evolution of the Genetic Code Revisited
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Genetic code: Lucky chance or fundamental law of nature?
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Origin and Evolutionary Process of the Genetic Code
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Flagged first, then by citations

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Sorted by citations

The N‐end rule pathway and regulation by proteolysis
Varshavsky, A. J. (2011) · Protein Science · doi:10.1002/pro.666 · Cited by 696 · see citing works
Codon usage bias
Parvathy, S. T., Udayasuriyan, V., & Bhadana, V. P. (2021) · Molecular Biology Reports · doi:10.1007/s11033-021-06749-4 · Cited by 579 · see citing works
Early bioenergetic evolution
Sousa, F. L., Thiergart, T., Landan, G., Nelson-Sathi, S., Pereira, I. A. C., Allen, J. F., Lane, N., & Martin, W. F. (2013) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2013.0088 · Cited by 286 · see citing works
“Given genes and proteins, the origin of molecular machines such as the ATPase is an admittedly impressive, but not conceptually challenging, evolutionary step; it is a far less problematic increment than the hurdles that had to be surmounted at the origin of the ribosome and the code [110,117].”
Background
Codon usage bias: causative factors, quantification methods and genome‐wide patterns: with emphasis on insect genomes
Behura, S. K., & Severson, D. W. (2012) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/j.1469-185x.2012.00242.x · Cited by 223 · see citing works
“As yet, the origin of the genetic code is still not fully understood (Koonin & Novozhilov, 2009), but its observed genomic pattern offers advances towards understanding the evolving nature of codon usage that is essential for the fidelity and efficiency of decoding genes to proteinsa process…”
BackgroundInfluential
Origin and Evolution of the Universal Genetic Code
Koonin, E. V., & Novozhilov, A. S. (2017) · Annual Review of Genetics · doi:10.1146/annurev-genet-120116-024713 · Cited by 200 · see citing works
Self-citation
RNA–Amino Acid Binding: A Stereochemical Era for the Genetic Code
Yarus, M., Widmann, J. J., & Knight, R. (2009) · Journal of Molecular Evolution · doi:10.1007/s00239-009-9270-1 · Cited by 198 · see citing works
“We accept the suggestion (Koonin and Novozhilov 2008) that random assignments in the manner of Crick’s ‘‘frozen accident’’ (Crick 1968) are also consistent with concurrent stereochemistry, co-evolution, and adaptation; all four together may have shaped the ultimate ‘universal’ genetic code.”
Background
Genetic Codes with No Dedicated Stop Codon: Context-Dependent Translation Termination
Swart, E. C., Serra, V., Petroni, G., & Nowacki, M. (2016) · Cell · doi:10.1016/j.cell.2016.06.020 · Cited by 167 · see citing works
“In light of the ambiguous genetic codes presented here, it is worth reconsidering the idea that the standard genetic code is ‘‘one in a million’’ and is optimized to minimize the effects of errors arising frommutations (Freeland and Hurst, 1998) (although contested [Koonin and Novozhilov, 2009]).”
Exoplanet Biosignatures: Future Directions
Walker, S. I., Bains, W., Cronin, L., DasSarma, S., Danielache, S. O., Domagal-Goldman, S. D., Kaçar, B., Kiang, N. Y.-L., Lenardic, A., Reinhard, C. T., et al. (2018) · Astrobiology · doi:10.1089/ast.2017.1738 · Cited by 122 · see citing works
“However, components of LUCA, such as archaeal and bacterial cell membrane components and metabolic capabilities, may have arisen independently multiple times.”
BackgroundInfluential
What is code biology?
Barbieri, M. (2017) · Biosystems · doi:10.1016/j.biosystems.2017.10.005 · Cited by 117 · see citing works
Modern diversification of the amino acid repertoire driven by oxygen
Granold, M., Hajieva, P., Toșa, M. I., Irimie, F. D., & Moosmann, B. (2017) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1717100115 · Cited by 86 · see citing works
Universality of the DNA methylation codes in Eucaryotes
Aliaga, B., Bulla, I., Mouahid, G., DUVAL, D. T., & Grunau, C. (2019) · Scientific Reports · doi:10.1038/s41598-018-37407-8 · Cited by 87 · see citing works
Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code
Koonin, E. (2017) · Life · doi:10.3390/life7020022 · Cited by 85 · see citing works
“, the mapping of 64 codons to 20 amino acids and the stop signal, is clearly non-random by multiple criteria [9,10].”
BackgroundInfluentialSelf-citation
Phenylalanine Increases Membrane Permeability
Perkins, R., & Vaida, V. (2017) · Journal of the American Chemical Society · doi:10.1021/jacs.7b09219 · Cited by 83 · see citing works
The Origin of Prebiotic Information System in the Peptide/RNA World: A Simulation Model of the Evolution of Translation and the Genetic Code
Chatterjee, S., & Yadav, S. (2019) · Life · doi:10.3390/life9010025 · Cited by 80 · see citing works
“<Table 1 about here> The origin of the genetic code remains elusive, even though the full codon catalog was deciphered over 50 years ago [93].”
Background
tRNA acceptor stem and anticodon bases form independent codes related to protein folding
Carter, C. W., & Wolfenden, R. V. (2015) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1507569112 · Cited by 79 · see citing works
Expanded use of sense codons is regulated by modified cytidines in tRNA
Cantara, W. A., Murphy, F. V., DeMi̇rci̇, H., & Agris, P. F. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1222641110 · Cited by 79 · see citing works
Inevitability of the emergence and persistence of genetic parasites caused by evolutionary instability of parasite-free states
Koonin, E. V., Wolf, Y. I., & Katsnelson, M. I. (2017) · Biology Direct · doi:10.1186/s13062-017-0202-5 · Cited by 74 · see citing works
“Namely, among different hypotheses, the stereochemical hypothesis of the code’s origin has received significant attention over the past decades [60, 61].”
BackgroundSelf-citation
A novel nuclear genetic code alteration in yeasts and the evolution of codon reassignment in eukaryotesPreprint
Mühlhausen, S., Findeisen, P., Plessmann, U., Urlaub, H., & Kollmar, M. (2016) · Genome Research · doi:10.1101/gr.200931.115 · Cited by 74 · see citing works
Refactored genetic codes enable bidirectional genetic isolation
Zürcher, J. F., Robertson, W. E., Kappes, T., Petris, G., Elliott, T. S., Salmond, G. P. C., & Chin, J. W. (2022) · Science · doi:10.1126/science.add8943 · Cited by 73 · see citing works
Ancestral Reconstruction of a Pre-LUCA Aminoacyl-tRNA Synthetase Ancestor Supports the Late Addition of Trp to the Genetic Code
Fournier, G. P., & Alm, E. J. (2015) · Journal of Molecular Evolution · doi:10.1007/s00239-015-9672-1 · Cited by 69 · see citing works
“2006; Wong 1988{Cavalcanti, 2004 #63)reviews in (Koonin and Novozhilov 2009; Trifonov 2000).”
Background
The cellular landscape of mid‐size noncoding RNA
Boivin, V., Faucher‐Giguère, L., Scott, M. S., & Elela, S. A. (2019) · Wiley Interdisciplinary Reviews - RNA · doi:10.1002/wrna.1530 · Cited by 70 · see citing works
“The discovery of tRNA and the standard genetic code added a whole new dimension to sequence selection in evolution (Itzkovitz & Alon, 2007; Koonin & Novozhilov, 2009; Pechmann & Frydman, 2013).”
Background
Insuperable problems of the genetic code initially emerging in an RNA world
Wills, P. R., & Carter, C. W. (2017) · Biosystems · doi:10.1016/j.biosystems.2017.09.006 · Cited by 64 · see citing works
Origins of Life: The Primal Self-Organization
Egel, R., Mulkidjanian, A. Y., & Lankenau, D. (2011) · Springer eBooks · doi:10.1007/978-3-642-21625-1 · Cited by 66 · see citing works
RNA ‐protein interactions in an unstructured context
Žagrović, B., Bartonek, L., & Polyansky, A. A. (2018) · FEBS Letters · doi:10.1002/1873-3468.13116 · Cited by 61 · see citing works
Short tRNA anticodon stem and mutant eRF1 allow stop codon reassignment
Kachale, A., Pavlíková, Z., Nenarokova, A., Roithová, A., Durante, I. M., Miletínová, P., Záhonová, K., Nenarokov, S., Votýpka, J., Horáková, E., et al. (2023) · Nature · doi:10.1038/s41586-022-05584-2 · Cited by 63 · see citing works
Interdependence, Reflexivity, Fidelity, Impedance Matching, and the Evolution of Genetic Coding
Carter, C. W., & Wills, P. R. (2017) · Molecular Biology and Evolution · doi:10.1093/molbev/msx265 · Cited by 61 · see citing works
Mutational properties of amino acid residues: implications for evolvability of phosphorylatable residues
Creixell, P., Schoof, E. M., Tan, C. S. H., & Linding, R. (2012) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2012.0076 · Cited by 59 · see citing works
Cracking the Code: Reprogramming the Genetic Script in Prokaryotes and Eukaryotes to Harness the Power of Noncanonical Amino Acids
Jann, C., Giofrè, S., Bhattacharjee, R., & Lemke, E. A. (2024) · Chemical Reviews · doi:10.1021/acs.chemrev.3c00878 · Cited by 54 · see citing works
Codon Size Reduction as the Origin of the Triplet Genetic Code
Baranov, P. V., Venin, M., & Provan, G. M. (2009) · PLoS ONE · doi:10.1371/journal.pone.0005708 · Cited by 55 · see citing works
“While the accidental nature of the genetic code is still a matter of scientific dispute [16,26–28], it is clear that the genetic code is”
Background
Frameshifting preserves key physicochemical properties of proteins
Bartonek, L., Braun, D., & Žagrović, B. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1911203117 · Cited by 53 · see citing works
Evidence of direct complementary interactions between messenger RNAs and their cognate proteins
Polyansky, A. A., & Žagrović, B. (2013) · Nucleic Acids Research · doi:10.1093/nar/gkt618 · Cited by 52 · see citing works
“As suggested before, it is possible that the answer lies in a combination of different influences (19).”
Beyond Terrestrial Biology: Charting the Chemical Universe of α-Amino Acid Structures
Meringer, M., Cleaves, H. J., & Freeland, S. J. (2013) · Journal of Chemical Information and Modeling · doi:10.1021/ci400209n · Cited by 53 · see citing works
Coding of Class I and II Aminoacyl-tRNA Synthetases
Carter, C. W. (2017) · Advances in experimental medicine and biology · doi:10.1007/5584_2017_93 · Cited by 52 · see citing works
“The most serious challenge to the various scenarios described in this review is the significant evidence from Koonin’s group (Koonin and Novozhilov 2009; Koonin 2011; Aravind et al. 2002; Leipe et al. 2002; Wolf et al. 1999) and others (Caetano-Anolles et al. 2007, 2013) that speciation of the…”
Background
Disruption of Transcriptional Coactivator Sub1 Leads to Genome-Wide Re-distribution of Clustered Mutations Induced by APOBEC in Active Yeast Genes
Lada, A. G., Kliver, S. F., Dhar, A., Polev, D. E., Masharsky, A., Rogozin, I. B., & Pavlov, Y. I. (2015) · PLoS Genetics · doi:10.1371/journal.pgen.1005217 · Cited by 52 · see citing works
“On the evolutionary scale, the general rule is that highly expressed genes are changing slowly (reviewed in [48, 49]).”
Background
Evolution of the genetic code: The ambiguity-reduction theory
Barbieri, M. (2019) · Biosystems · doi:10.1016/j.biosystems.2019.104024 · Cited by 53 · see citing works
Ribosome Structure, Function, and Early Evolution
Opron, K., & Burton, Z. F. (2018) · International Journal of Molecular Sciences · doi:10.3390/ijms20010040 · Cited by 50 · see citing works
FACIL: Fast and Accurate Genetic Code Inference and Logo
Dutilh, B. E., Jurgelenaite, R., Szklarczyk, R., van Hijum, S. A. F. T., Harhangi, H. R., Schmid, M. C., de Wild, B., Françoijs, K., Stunnenberg, H. G., Strous, M., et al. (2011) · Bioinformatics · doi:10.1093/bioinformatics/btr316 · Cited by 49 · see citing works
“After all the genetic code or codon table, i.e. the ‘dictionary’ that translates codons (nucleic acid triplets) ∗To whom correspondence should be addressed. into amino acids (AAs), is largely universal and unambiguous (Koonin and Novozhilov, 2009).”
BackgroundMethod
Bacterial Diversification in the Light of the Interactions with Phages: The Genetic Symbionts and Their Role in Ecological Speciation
Braga, L. P. P., Soucy, S. M., Amgarten, D., da Silva, A. M., & Setúbal, J. C. (2018) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2018.00006 · Cited by 49 · see citing works
“Koonin and Novozhilov (2009) have pointed out the relation between HGT and the universality of the genetic code, allowing the flow of information necessary for evolution among biological entities in all directions (i.e., vertically and horizontally).”
Background
Optimization of the standard genetic code according to three codon positions using an evolutionary algorithm
Błażej, P., Wnętrzak, M., Mackiewicz, D., & Mackiewicz, P. (2018) · PLoS ONE · doi:10.1371/journal.pone.0201715 · Cited by 51 · see citing works
Evolution of Life on Earth: tRNA, Aminoacyl-tRNA Synthetases and the Genetic Code
Lei, L., & Burton, Z. F. (2020) · Life · doi:10.3390/life10030021 · Cited by 50 · see citing works
“Figure 6 shows the 4-amino acid code to encode Gly, Asp, Ala, and Val [1,6,46,47].”
Background
Sequence signatures of direct complementarity between mRNAs and cognate proteins on multiple levels
Hlevnjak, M., Polyansky, A. A., & Žagrović, B. (2012) · Nucleic Acids Research · doi:10.1093/nar/gks679 · Cited by 44 · see citing works
“Previous computational analyses have shown that the genetic code is robust to random point mutations and translational errors, such that hydrophobicity of the original and the mutated amino acids differs to a minimal degree (‘error-minimization’ hypothesis) (2,13,43).”
BackgroundInfluential
Synthetic transitions: towards a new synthesis
Solé, R. V. (2016) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2015.0438 · Cited by 42 · see citing works
Universal biology and the statistical mechanics of early life
Goldenfeld, N. D., Biancalani, T., & Jafarpour, F. (2017) · Philosophical Transactions of the Royal Society A Mathematical Physical and Engineering Sciences · doi:10.1098/rsta.2016.0341 · Cited by 46 · see citing works
“system models to study two of the most perplexing features shared by all known life on the Earth: (i) the canonical genetic code [8] and (ii) biological homochirality [9].”
Background
Absolute binding-free energies between standard RNA/DNA nucleobases and amino-acid sidechain analogs in different environments
de Ruiter, A., & Žagrović, B. (2014) · Nucleic Acids Research · doi:10.1093/nar/gku1344 · Cited by 43 · see citing works
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Frozen Accident Pushing 50: Stereochemistry, Expansion and Chance in the Evolution of the Genetic Code
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Authors: Koonin, E. V., & Novozhilov, A. S.
Affiliations: National Institutes of Health (US)
Corresponding author: Koonin, Eugene V.
ORCID: Koonin, Eugene V.
Novozhilov, Artem S.
Venue: IUBMB Life
Publisher: Wiley
Published: 2008-12-31
Volume: 61
Issue: 2
Pages: 99-111
ISSN: 1521-6543 (ISSN-L)
1521-6551
DOI: 10.1002/iub.146
PMC: PMC3293468
OpenAlex: W1963573192
Cited by: 392
References: 143
Citations by year: 2026: 36 · 2025: 22 · 2024: 28 · 2023: 33 · 2022: 21 · 2021: 21 · 2020: 28 · 2019: 22 · 2018: 25 · 2017: 32 · 2016: 18 · 2015: 19 · 2014: 14 · 2013: 20 · 2012: 16
FWCI: 0.95
Type: journal article
Language: en
Open access: green
www.ncbi.nlm.nih.gov
Licence: onlinelibrary.wiley.com/termsAndConditions#vor (vor)
Locations: IUBMB Life (publishedVersion)
PubMed (publishedVersion)
Europe PMC (PubMed Central) (acceptedVersion)
PubMed Central (acceptedVersion)
Funders: Aspen Center for Physics
National Institutes of Health
U.S. National Library of Medicine (ZIALM000061)
Topics: Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › RNA and protein synthesis mechanisms
Physical Sciences › Physics and Astronomy › Astronomy and Astrophysics › Origins and Evolution of Life
Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Evolution and Genetic Dynamics
Keywords: origin of the genetic code, genetic code evolution, codon assignment, error reduction, codon reassignment, code robustness
MeSH terms: Amino Acids, Codon, Genetic Code, Protein Biosynthesis, Evolution, Molecular

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Origin and evolution of the genetic code: The universal enigma

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