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A primordial genetic code is proposed, having only four codons assigned, GGCmeaning glycine, GAC meaning aspartate/glutamate, GCC meaning alanine-like andGUC meaning valine-like. Pathways of ambiguity reduction enlarged the codonrepertoire with CUC meaning leucine, AUC meaning isoleucine, ACC meaningthreonine-like and GAG meaning glutamate. Introduction of UNN anticodons, in anext episode of code evolution in which nonsense elimination was the leadingtheme, introduced a family box structure superposed on the original mirrorstructure. Finally, growth rate was the leading theme during the remainingrepertoire expansion, explaining the ordered phylogenetic pattern ofaminoacyl-tRNA synthetases. The special role of natural aptamers in the processis high-lighted, and the error robustness characteristics of the code are shownto have evolved by way of a stepwise, restricted enlargement of the tRNArepertoire, instead of by an exhaustive selection process testing myriads ofcodes.

Three phases in the evolution of the standard genetic code: how translation could get started