The most striking difference between prokaryotes and eukaryotes is the
way in which their genetic material is packaged. Eukaryotic cells have two or more chromosomes contained within a membrane-delimited organelle, the nucleus. In contrast, prokaryotes lack a membrane- delimited nucleus. The prokaryotic chromosome is located in an irregularly shaped region called the nucleoid (other names are also used: the nuclear body, chromatin body, nuclear region) (figure 3.16). Usually prokaryotes contain a single circle of double-stranded deoxyribonucleic acid (DNA), but some have a linear DNA chromosome and some, such as Vibrio cholerae and Borrelia burgdorferi (the causative agents of cholera and Lyme disease, respectively), have more than one chromosome. Both electron and light microscopic studies have been important for understanding nucleoid structure and function, especially during active cell growth and division. The nucleoid has a fibrous appearance in electron micrographs; the fibers are probably DNA. In actively growing cells, the nucleoid has projections that extend into the cytoplasmic matrix. Presumably these projections contain DNA that is being actively transcribed to produce mRNA. Other studies have shown that more than one nucleoid can be observed within a single cell when genetic material has been duplicated but cell division has not yet occurred (figure 3.16a). It is possible to isolate pure nucleoids. Chemical analysis of purified nucleoids reveals that they are composed of about 60% DNA, 30% RNA, and 10% protein by weight. In Escherichia coli, the closed DNA circle measures approximately 1,400 um or about 230–700 times longer than the cell (figure 3.16b). Obviously it must be very efficiently packaged to fit within the nucleoid. The DNA is looped and coiled extensively, probably with the aid of RNA and a variety of nucleoid proteins. These include condensing proteins, which are conserved in both Bacteria and Archaea. Unlike the eukaryotes and some archaea, Bacteria do not use histone proteins to package their DNA. There are a few exceptions to the preceding picture. Membrane-bound DNA-containing regions are present in two genera of the unusual bacterial phylum Planctomycetes (see figure 21.12). Pirellula has a single membrane that surrounds a region, the pirellulosome, which contains a fibrillar nucleoid and ribosome- like particles. The nuclear body of Gemmata obscuriglobus is bounded by two membranes. More work will be required to determine the functions of these membranes and how widespread this phenomenon is.