Table 2.

The Rates of Synonymous (Ks) and Nonsynonymous (Ka) Nucleotide Substitutions among Different Functional Categories for Essential, Nonessential, and Undetermined Escherichia coliGenes

Ks (±se)[i] Ka (±se)[ii] Ka/Ks (±se)
Information storage and processing
 Essential (n = 113)19.42 × 10−3 0.96 × 10−3 3.89 × 10−2
(±2.3 × 10−3)(±0.2 × 10−3)(±0.8 × 10−2)
 Nonessential (n = 222)48.16 × 10−3 3.50 × 10−3 8.72 × 10−2
(±2.8 × 10−3)(±0.4 × 10−3)(±1.1 × 10−2)
 Undetermined (n = 113)52.31 × 10−3 4.62 × 10−3 11.69 × 10−2
(±4.4 × 10−3)(±0.6 × 10−3)(±1.8 × 10−2)
 Significance of the difference[iii] essential vs. nonessential P = 1.6 × 10−15 P = 1.6 × 10−7 P = 6.1 × 10−3
 Significance of the difference[iii] essential vs. undetermined P = 3.2 × 10−12 P = 1.3 × 10−10 P = 1.3 × 10−4
 Significance of the difference[iii] nonessential vs. undeterminedNSNSNS
Cellular processes
 Essential (n = 46)37.57 × 10−3 1.32 × 10−3 4.72 × 10−2
(±3.5 × 10−3)(±0.3 × 10−3)(±1.5 × 10−2)
 Nonessential (n = 408)47.17 × 10−3 3.01 × 10−3 7.86 × 10−2
(±1.9 × 10−3)(±0.3 × 10−3)(±0.7 × 10−2)
 Undetermined (n = 154)40.92 × 10−3 4.10 × 10−3 8.80 × 10−2
(±2.1 × 10−3)(±0.5 × 10−3)(±0.9 × 10−2)
 Significance of the difference[iii] essential vs. nonessentialNS P = 3.6 × 10−2 NS
 Significance of the difference[iii] essential vs. undeterminedNS P = 2.0 × 10−4 P = 3.7 × 10−4
 Significance of the difference[iii] nonessential vs. undeterminedNS P = 2.8 × 10−3 P = 3.2 × 10−4
Metabolism
 Essential (n = 34)37.18 × 10−3 1.09 × 10−3 3.19 × 10−2
(±8.1 × 10−3)(±0.3 × 10−3)(±1.1 × 10−2)
 Nonessential (n = 725)54.76 × 10−3 2.52 × 10−3 5.92 × 10−2
(±1.5 × 10−3)(±0.1 × 10−3)(±0.4 × 10−2)
 Undetermined (n = 241)59.32 × 10−3 4.50 × 10−3 9.80 × 10−2
(±3.2 × 10−3)(±0.3 × 10−3)(±0.8 × 10−2)
 Significance of the difference[iii] essential vs. nonessential P = 1.2 × 10−4 P = 1.2 × 10−3 P = 2.7 × 10−3
 Significance of the difference[iii] essential vs. undetermined P = 8.2 × 10−5 P = 4.9 × 10−8 P = 1.7 × 10−6
 Significance of the difference[iii] nonessential vs. undeterminedNS P = 3.6 × 10−13 P = 8.2 × 10−11
Poorly Characterized
 Essential (n = 12)28.75 × 10−3 1.75 × 10−3 11.20 × 10−2
(±6.0 × 10−3)(±0.3 × 10−3)(±4.2 × 10−2)
 Nonessential (n = 439)49.59 × 10−3 5.93 × 10−3 13.08 × 10−2
(±2.1 × 10−3)(±0.6 × 10−3)(±0.9 × 10−2)
 Undetermined (n = 599)45.51 × 10−3 5.51 × 10−3 12.60 × 10−2
(±1.7 × 10−3)(±0.5 × 10−3)(±0.8 × 10−2)
 Significance of the difference[iii] essential vs. nonessentialNSNSNS
 Significance of the difference[iii] essential vs. undeterminedNSNSNS
 Significance of the difference[iii] nonessential vs. undeterminedNSNSNS

[i] The average synonymous substitution (nucleotide substitutions that do not change the encoded amino acid sequence) rate (Ks) for all orthologous genes within a given fitness class and species is shown with the standard error (in parentheses).

[ii] The average nonsynonymous substitution (nucleotide substitutions that change the encoded amino acid sequence) rate (Ka) for all orthologous genes within a given fitness class and species is shown with the standard error (in parentheses).

[iii] Statistical significance of the difference between the essential and nonessential classes for a given measurement as determined using the Mann-Whitney U test.