  The genetic code has a high level of error robustness. Using values of
hydrophobicity scales as a proxy for amino acid character, and the Mean Square
measure as a function quantifying error robustness, a value can be obtained for
a genetic code which reflects the error robustness of that code. By comparing
this value with a distribution of values belonging to codes generated by random
permutations of amino acid assignments, the level of error robustness of a
genetic code can be quantified. We present a calculation in which the standard
genetic code is shown to be optimal. We obtain this result by (1) using
recently updated values of polar requirement as input; (2) fixing seven
assignments (Ile, Trp, His, Phe, Tyr, Arg, and Leu) based on aptamer
considerations; and (3) using known biosynthetic relations of the 20 amino
acids. This last point is reflected in an approach of subdivision (restricting
the random reallocation of assignments to amino acid subgroups, the set of 20
being divided in four such subgroups). The three approaches to explain
robustness of the code (specific selection for robustness, amino acid-RNA
interactions leading to assignments, or a slow growth process of assignment
patterns) are reexamined in light of our findings. We offer a comprehensive
hypothesis, stressing the importance of biosynthetic relations, with the code
evolving from an early stage with just glycine and alanine, via intermediate
stages, towards 64 codons carrying todays meaning.
