Rewiring the keyboard evolvability of the genetic code

The genetic code evolved in two distinct phases. First, the 'canonical' code emerged before the last universal ancestor; subsequently this code diverged in numerous nuclear and organelle lineages. Here, we examine the distribution and causes of these secondary deviations from the canonical genetic code. The majority of non-standard codes arise from alterations in the tRNA, with most occurring by post-transcriptional modifications, such as base modification or RNA editing, rather than by substitutions within tRNA anticodons.