The general characteristics of the reference genomes are summarized in Table 3. The genome size ranged from 2 167 361 to 2 292 467 bp. The number of coding sequences (CDSs) after manual curation varied from 2295 to 2450 with an average CDS size of 796.6 bp. The number of core genes was 1820 and accessory genes ranged from 475 to 630. In Figure 1(a and b) BLAST atlases of the 14 reference genomes and all identified plasmid sequences, respectively, are displayed. Briefly, the reference genomes showed relatively high genomic similarity among all strains with the exception of two large insertions in WHO F and the presence or absence of the gonococcal genomic island (GGI).58 The insertions in WHO F each include 34 almost identical predicted CDSs (31 984 bp) mainly containing an apparently complete type IV secretion system, a vapD virulence gene and xerC recombinase inserted into a tRNA-Asn element, with only weak matches to other sequences in the NCBI non-redundant nt database. Other regions with low genetic conservation corresponded to mobile genetic elements and prophages (Figure 1a).
Table 3.

General characteristics of the reference genomes of the 2016 WHO N. gonorrhoeae reference strains (n = 14)

CharacteristicWHO FWHO GWHO KWHO LWHO MWHO NWHO OWHO PWHO UWHO VWHO WWHO XWHO YWHO Z
Genome size (bp)2 292 4672 167 3612 169 8462 168 6332 178 3442 172 8262 169 0622 173 8612 234 2692 221 2842 222 3862 171 1122 228 9802 229 351
No. of CDSs24502299229623142305230023042305237823662361229523802368
Coding density (%)84.884.584.684.584.584.684.684.684.884.884.784.584.884.6
Average gene size (bp)793.7796.5799.3791.6798.7798.9796.0797.6796.8795.8797.5799.5794.4796.7
GC content (%)52.152.652.652.652.652.652.652.652.452.452.452.652.452.4
5S rRNA4
16S rRNA4
23S rRNA4
tRNAs5656575656565656565656575657
ncRNAs1615151515151616161616161616
tmRNA11111111111111
No. of genes in pangenome3478
No. core genes1820
Accessory genes (%)630 (25.7)479 (20.8)476 (20.7)494 (21.3)485 (21.0)480 (20.9)484 (21.0)485 (21.0)558 (23.5)546 (23.1)541 (22.9)475 (20.7)560 (23.5)548 (23.1)
DNA uptake sequences (DUSs)a1981 (1533)1947 (1510)1950 (1510)1956 (1518)1955 (1516)1951 (1513)1950 (1519)1959 (1517)1963 (1512)1968 (1518)1954 (1509)1949 (1510)1973 (1522)1959 (1512)
Number of plasmids02123331122111
CharacteristicWHO FWHO GWHO KWHO LWHO MWHO NWHO OWHO PWHO UWHO VWHO WWHO XWHO YWHO Z
Genome size (bp)2 292 4672 167 3612 169 8462 168 6332 178 3442 172 8262 169 0622 173 8612 234 2692 221 2842 222 3862 171 1122 228 9802 229 351
No. of CDSs24502299229623142305230023042305237823662361229523802368
Coding density (%)84.884.584.684.584.584.684.684.684.884.884.784.584.884.6
Average gene size (bp)793.7796.5799.3791.6798.7798.9796.0797.6796.8795.8797.5799.5794.4796.7
GC content (%)52.152.652.652.652.652.652.652.652.452.452.452.652.452.4
5S rRNA4
16S rRNA4
23S rRNA4
tRNAs5656575656565656565656575657
ncRNAs1615151515151616161616161616
tmRNA11111111111111
No. of genes in pangenome3478
No. core genes1820
Accessory genes (%)630 (25.7)479 (20.8)476 (20.7)494 (21.3)485 (21.0)480 (20.9)484 (21.0)485 (21.0)558 (23.5)546 (23.1)541 (22.9)475 (20.7)560 (23.5)548 (23.1)
DNA uptake sequences (DUSs)a1981 (1533)1947 (1510)1950 (1510)1956 (1518)1955 (1516)1951 (1513)1950 (1519)1959 (1517)1963 (1512)1968 (1518)1954 (1509)1949 (1510)1973 (1522)1959 (1512)
Number of plasmids02123331122111

aTotal of the 10-mer DUS sequence GCCGTCTGAA (no. of the 12-mer ATGCCGTCTGAA). Note: the 10-mer sequence is included in the 12-mer.

Table 3.

General characteristics of the reference genomes of the 2016 WHO N. gonorrhoeae reference strains (n = 14)

CharacteristicWHO FWHO GWHO KWHO LWHO MWHO NWHO OWHO PWHO UWHO VWHO WWHO XWHO YWHO Z
Genome size (bp)2 292 4672 167 3612 169 8462 168 6332 178 3442 172 8262 169 0622 173 8612 234 2692 221 2842 222 3862 171 1122 228 9802 229 351
No. of CDSs24502299229623142305230023042305237823662361229523802368
Coding density (%)84.884.584.684.584.584.684.684.684.884.884.784.584.884.6
Average gene size (bp)793.7796.5799.3791.6798.7798.9796.0797.6796.8795.8797.5799.5794.4796.7
GC content (%)52.152.652.652.652.652.652.652.652.452.452.452.652.452.4
5S rRNA4
16S rRNA4
23S rRNA4
tRNAs5656575656565656565656575657
ncRNAs1615151515151616161616161616
tmRNA11111111111111
No. of genes in pangenome3478
No. core genes1820
Accessory genes (%)630 (25.7)479 (20.8)476 (20.7)494 (21.3)485 (21.0)480 (20.9)484 (21.0)485 (21.0)558 (23.5)546 (23.1)541 (22.9)475 (20.7)560 (23.5)548 (23.1)
DNA uptake sequences (DUSs)a1981 (1533)1947 (1510)1950 (1510)1956 (1518)1955 (1516)1951 (1513)1950 (1519)1959 (1517)1963 (1512)1968 (1518)1954 (1509)1949 (1510)1973 (1522)1959 (1512)
Number of plasmids02123331122111
CharacteristicWHO FWHO GWHO KWHO LWHO MWHO NWHO OWHO PWHO UWHO VWHO WWHO XWHO YWHO Z
Genome size (bp)2 292 4672 167 3612 169 8462 168 6332 178 3442 172 8262 169 0622 173 8612 234 2692 221 2842 222 3862 171 1122 228 9802 229 351
No. of CDSs24502299229623142305230023042305237823662361229523802368
Coding density (%)84.884.584.684.584.584.684.684.684.884.884.784.584.884.6
Average gene size (bp)793.7796.5799.3791.6798.7798.9796.0797.6796.8795.8797.5799.5794.4796.7
GC content (%)52.152.652.652.652.652.652.652.652.452.452.452.652.452.4
5S rRNA4
16S rRNA4
23S rRNA4
tRNAs5656575656565656565656575657
ncRNAs1615151515151616161616161616
tmRNA11111111111111
No. of genes in pangenome3478
No. core genes1820
Accessory genes (%)630 (25.7)479 (20.8)476 (20.7)494 (21.3)485 (21.0)480 (20.9)484 (21.0)485 (21.0)558 (23.5)546 (23.1)541 (22.9)475 (20.7)560 (23.5)548 (23.1)
DNA uptake sequences (DUSs)a1981 (1533)1947 (1510)1950 (1510)1956 (1518)1955 (1516)1951 (1513)1950 (1519)1959 (1517)1963 (1512)1968 (1518)1954 (1509)1949 (1510)1973 (1522)1959 (1512)
Number of plasmids02123331122111

aTotal of the 10-mer DUS sequence GCCGTCTGAA (no. of the 12-mer ATGCCGTCTGAA). Note: the 10-mer sequence is included in the 12-mer.

BLAST atlas of the 2016 WHO N. gonorrhoeae reference genomes (n = 14). (a) A genome comparison of the 2016 WHO reference strains presented in this study using WHO F23 as reference and (b) a comparison of the up to three plasmids (named pCryptic, pBlaTEM and pConjugative65,81,82) identified in the same strains. WHO G pCryptic (cryptic plasmid), WHO M pBlaTEM (β-lactamase-producing plasmid) and WHO G pConjugative [conjugative plasmid including tet(M) in WHO G and N] were used as references, respectively. For each, GC skew is shown in the inner rings. The position in the genomes is shown for genetic resistance determinants and loci used for molecular diagnostics and epidemiological characterization, i.e. NG-MAST STs and MLST STs. The presence of the GGI58 is indicated in red. An approximately 500 bp region with lower nucleotide conservation (∼75% identity) is shown with lighter colours in WHO M and WHO O pBlaTEM plasmids corresponding to a hypothetical protein.
Figure 1.

BLAST atlas of the 2016 WHO N. gonorrhoeae reference genomes (n = 14). (a) A genome comparison of the 2016 WHO reference strains presented in this study using WHO F23 as reference and (b) a comparison of the up to three plasmids (named pCryptic, pBlaTEM and pConjugative65,81,82) identified in the same strains. WHO G pCryptic (cryptic plasmid), WHO M pBlaTEM (β-lactamase-producing plasmid) and WHO G pConjugative [conjugative plasmid including tet(M) in WHO G and N] were used as references, respectively. For each, GC skew is shown in the inner rings. The position in the genomes is shown for genetic resistance determinants and loci used for molecular diagnostics and epidemiological characterization, i.e. NG-MAST STs and MLST STs. The presence of the GGI58 is indicated in red. An approximately 500 bp region with lower nucleotide conservation (∼75% identity) is shown with lighter colours in WHO M and WHO O pBlaTEM plasmids corresponding to a hypothetical protein.

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