Table 1.

The Rates of Synonymous (Ka) and Nonsynonymous (Ks) Nucleotide Substitutions for Essential Versus Nonessential Bacterial Genes

Ks (±se)[i] Ka (±se)[ii] r[iii] Ka/Ks
Escherichia coli—all[iv] 26.99 × 10−3 1.11 × 10−3 0.354.50 × 10−2
 Essential (n = 205)(±2.1 × 10−3)(±0.1 × 10−3)(±0.7 × 10−2)
 Nonessential (n = 1794)51.0 × 10−3 3.60 × 10−3 0.448.40 × 10−2
(±1.0 × 10−3)(±0.2 × 10−3)(±0.2 × 10−2)
 Undetermined (n = 1107)48.6 × 10−3 5.00 × 10−3 0.4711.67 × 10−2
(±1.3 × 10−3)(±0.3 × 10−3)(±0.5 × 10−2)
 Significance of the difference[vi] P = 0 P = 5.6 × 10−16 na P = 6.4 × 10−6
E. coli—experimental[v] 35.33 × 10−3 1.36 × 10−3 0.334.73 × 10−6
 Essential (n = 150)(±2.5 × 10−3) (±0.2 × 10−3)(±0.7 × 10−2)
 Nonessential (n = 1736)51.29 × 10−3 3.60 × 10−3 0.448.27 × 10−2
(±1.0 × 10−3)(±0.2 × 10−3)(±0.4 × 10−2)
 Significance of the difference[vi] P = 1.2 × 10−7 P = 1.2 × 10−8 na P = 5.6 × 10−4
Helicobacter pylori111.33 × 10−3 12.89 × 10−3 0.4311.32 × 10−2
 Essential (n = 98)(±4.1 × 10−3)(±1.1 × 10−3)(±0.9 × 10−2)
 Nonessential (n = 130)135.24 × 10−3 21.64 × 10−3 0.3016.14 × 10−2
(±3.1 × 10−2)(±1.4 × 10−3)(±0.8 × 10−2)
 Significance of the difference[vi] P = 3.5 × 10−6 P = 6.1 × 10−7 na P = 6.1 × 10−4
Neisseria meningitidis65.37 × 10−3 4.76 × 10−3 0.687.32 × 10−2
 Essential (n = 98)(±6.9 × 10−3)(±0.7 × 10−3)(±1.0 × 10−2)
 Nonessentials (n = 130)91.56 × 10−3 9.60 × 10−3 0.4917.65 × 10−2
(±5.9 × 10−3)(±0.9 × 10−3 (±1.9 × 10−2)
 Significance of the difference[vi] P = 1.6 × 10−4 P = 1.0 × 10−7 na P = 4.2 × 10−5

[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] Correlation coefficient between the Ks and Ka values.

[iv] This set includes all of the E. coli genes characterized as essential or nonessential (on the basis of experimental and functional information).

[v] This set includes only the E. coli genes characterized as essential or nonessential on the basis of experimental evidence.

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