Frameshift and wild-type proteins are often highly similar because the genetic code and genomes were optimized for frameshift tolerance

Table 7 The usage of codon pairs and their weighted mean FSSs (Gon250)

No	species	Number of codon pairs			Weighted mean FSS
No	species	Over-represented	Under-represented	Absent	Over-represented	Under-represented	All
1	H. sapiens	1573	2523	50	−1.52	−7.80	−3.06
2	M. musculus	1505	2591	190	−2.83	−7.13	− 3.81
3	X. tropicalis	1660	2436	148	−3.12	−6.98	−3.80
4	D. rerio	1493	2603	148	−4.87	−6.09	−5.18
5	D. melanogaster	1418	2678	140	−5.33	−5.86	−5.02
6	C. elegans	1469	2627	164	−6.47	−5.26	−6.11
7	A. thaliana	1566	2530	15	−6.30	−5.35	− 6.37
8	S. cerevisiae	1493	2603	159	−4.86	−6.14	−4.27
9	E. coli	1389	2707	197	−6.76	− 5.11	− 6.82
10	Equal Usage	0	0	0	N/A	N/A	−5.67

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