The literature

Every source behind the genetic-code optimality article, what it measured, whether anyone here opened it, which way it cuts, and the sources considered and not used.

The larger catalogue for the whole series is on the parent page, the origin of life:  the literature.

Sources cited

Source What it is Access Direction
Wong 1975, PNAS 72:1909 The coevolution theory, on eight precursor-product pairs. Cited through Koonin & Novozhilov 2009 complicates
Freeland & Hurst 1998, J Mol Evol 47:238 "The genetic code is one in a million," within a space of block-preserving shuffles. Abstract only (Springer paywall) complicates
Novozhilov, Wolf & Koonin 2007, Biol Direct 2:24 The greedy-optimisation computation behind the swap-count and mediocre-peak results. Opened in full (CC BY) undercuts the optimality argument
Koonin & Novozhilov 2009, IUBMB Life 61:99 The review that states the two-trillion consequence, the 70-78% minimisation, the 10-12 swaps, the "not locally stable" finding, the cost-function caveat, and the Amirnovin recount. Opened in full (PMC author manuscript) complicates
Koonin 2017, Life 7:22 The 10-7-to-10-8 figure with the 1084 denominator in the same sentence, the methodological warning, and the fifty-year verdict on codon assignments. Opened in full (CC BY) supports the unsolved claim
Shulgina & Eddy 2021, eLife 10:e71402 251,571 genomes screened; five new reassignments; the codon-capture route. Opened in full (CC BY) undercuts universality
DeBenedictis, Söll & Esvelt 2022, Nucleic Acids Res 50:5906 All twenty transfer RNA isoacceptor classes converted to quadruplets; only nine synthetases tolerate them. Opened (CC BY) undercuts "only triplets work"
McGowan et al. 2023 A ciliate decoding UAA and UAG as two different amino acids. Opened (CC BY) undercuts universality
Wehbi et al. 2024, PNAS 121:e2410311121 The recruitment order re-derived from ancestral sequences, and the Miller-Urey sampling bias in the textbook order. Opened (CC BY) complicates
Nyerges et al. 2026 Recoded E. coli: "synonymous codon replacement — recoding — is frequently lethal." Post-cutoff. Opened complicates
Miyachi & Ichihashi 2026, eLife 15:RP111164 The first experimental test of error minimization. Prediction not detected; standard code the multi-property cost optimum among 19,683. Post-cutoff. Opened in full (CC BY) complicates, both ways
Hashmi et al. 2026 A converging amino acid chronology from sixteen molecular-complexity metrics. Post-cutoff. Opened (CC BY) complicates
Goldman, Fournier & Kaçar 2026, Cell Genomics 6:101140 Reconstructed pre-LUCA synthetases lacking tryptophan, and an ancestor that could not tell leucine from isoleucine from valine. Post-cutoff. Opened complicates

Considered and not used

  • Freeland & Hurst 1998 body. Springer paywall. Only the abstract was read. The description of the sampled space comes from Koonin and Novozhilov's account of it, which was opened.
  • Freeland et al. 2000, Mol Biol Evol 17:511. The source of the stronger "at or very close to a global optimum" claim. Not opened (Oxford University Press blocks automated access). This project's notes carry a "96% and 100%" figure from its body that could not be verified, and the article prints neither number. Printing 96-100% beside the 70-78% minimisation percentage would be a factual error, because they are different measures on different constraint sets.
  • Crick 1968, the frozen-accident paper. Named as a position, not quoted.
  • Amirnovin 1997. The recount of the coevolution theory. Not opened; Koonin's report of the three figures is what the article cites, and the footnote says so.
  • Ronneberg, Landweber & Freeland's wobble recount, which Koonin says removes the statistical support for coevolution altogether. Not opened; mentioned in the footnote rather than argued from.
  • Worning & Ibarra-Chávez 2026, on the twenty-seven NCBI translation tables and near-universality as a horizontal-transfer effect. Not opened; the table count is used and the interpretation is not.
  • Koonin 2014's "ominous specter of irreducible complexity" passage. It belongs to the translation article, not to the optimality question, and is used there.

Open questions

  1. Freeland et al. 2000 should be obtained. It is the source of the strongest optimality claim in circulation and nobody on this project has read it.
  2. Whether the Miyachi and Ichihashi result holds with more than three reassigned amino acids, and with proteins other than the three reporters used.
  3. Every figure in this article depends on a cost function chosen by the analyst. Koonin's own caveat about that is quoted; nobody appears to have tested how the rankings move under a genuinely different measure of amino acid similarity.
  4. An attribution error in this project's own dossier was found and corrected here. The "10–12 evolutionary steps" and "not locally stable" sentences were credited to Novozhilov, Wolf and Koonin 2007; they are in the 2009 review. Any other article quoting them should be checked.