Prokaryotes

Prokaryotic genomes are composed of a chromosome plus various accessory elements. The former is most commonly a circular double-stranded DNA molecule but may be a linear molecule in some major groups, such as Streptomyces and Borrelia (the causative agent of Lyme disease). Acces-

?sory elements most prominently include plasmids (commonly circular but linear in Actinomycetes and some Proteobacteria) as well as insertion sequence (IS) elements, transposons, and prophages (derived from f viruses). Other variations in chromosomal geometry exist: multiple cir cular chromosomes are found in some organisms; combinations of circular and linear chromosomes occur in others; and, in the extreme (observed in Streptomyces), circular and linear chromosomes can convert between those two topologies.

The smallest bacterial chromosome, with only 580 kilobase pairs (kbp) occurs in Mycoplasma genitalium, and the largest, with 9,200 kbp, occurs in Myxococcusxanthus. Representative sizes cluster between 2,000 and 5,000 kbp (e.g., Escherichia coli MG1655 has 4,649,221 bp). A typical bacterial gene contains approximately a thousand base pairs. M. genitalium has approximately 470 genes, while M. xanthus has more than 10,000, and E. coli has approximately 4,288.

By 2002 the nucleotide sequences of more than seventy-five prokaryotic chromosomes had been mapped. One goal of these sequencing projects allelic variation pres- is gene annotation: establishing the location, function, and allelic variation ence of different gene for each gene. In E. coli MG1655, for example, the positions of the 4,288 forms (alleles) in a . .. . . . . .., . . .. .

protein-coding genes have been identified; the average distance between population genes is 118 base pairs; and the noncoding sequences (some of which may function as regulatory sites) constitute less than 11 percent of the genome. The function of approximately 40 percent of the genes, however, remains unknown. Notably, the chromosomal size and gene content of another isolate of E. coli, the pathogenic H157:O7 strain, are quite different. The H157:O7 chromosome is 20 percent larger, while MG1655 and H157:O7 share 4.1 million base pairs (mbp) in common. H157:O7 has 1.34 mbp that are not found in MG1655 and MG1655 has 0.53 mbp that are not found in H157:O7.

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The genomes of closely related prokaryotes often have different organizations. These differences arise from rearrangements (such as inversions) between repeated elements, IS elements, and transposons and from the "horizontal transfer" of nucleotide sequences between cells. The latter phenomenon is mediated most commonly by conjugative plasmids, which are nonessential, autonomous accessory genetic elements that can acquire genes (such as antibiotic resistance genes) and then move them from a donor organism to a recipient. The dynamic character of genomic organization in prokaryotes is often designated as "genomic plasticity."

A series of repeated elements exist in the chromosomes of prokaryotes. In some instances the repeats are redundant copies of essential, long nucleotide sequences, as is seen in ribosomal RNA loci. Other repeats are small and have known functions (as in the Chi sequences in E. coli that facilitate genetic crossing over) or unknown functions (as in the REP [repeated extragenic palindromic] sequences in E. coli).

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