The genetic code is degenerate, but alternative synonymous codons are generally not used with equal frequency. Since the pioneering work of Grantham'a group (1, 2) it has been apparent that genes from one species often share similarities incodon frequency; under the “genome hypothesis” (1, 2) there is a species-specific pattern to codon usage.

However, it has become clear that in most species there are also considerable differences among genes (3–7). Multivariate analyses have revealed that in each species so far examined there is a single major trend in codon usage among genes, usually from highly biased to more nearly even usage of synonymous codons. Thus, to represent the codon usage pattern of an organism it is not sufficient to sum over all genes (8), as this conceals the underlying heterogeneity. Rather, it is necessary to describe the trend among genes seen in that species. We illustrate these trends for six species where codon usage has been examined in detail, by presenting the pooled codon usage for the 10% of genes at either end of the major trend (Table 1).

Closely-related organisms have similar patterns of codon usage, and so the six species in Table 1 are representative of wider groups. For example, with respect to codon usage, Salmonella typhimurlum closely resembles E. coli (9), while all mammalian species so far examined (principally mouse, rat and cow) largely resemble humans (4, 8).