Procaryotic Domains

Single cells or simple associations of similar cells (usually 0.210.0 m in smallest dimension, although some are much larger) forming a group defined by cellular, not organismal, properties (i.e., by the structure and components of the cells of an organism rather than by the properties of the organism as a whole)

The nucleoplasm (genophore) is, with a few exceptions, not separated from the cytoplasm by a unit-membrane system (nuclear membrane). Cell division is not accompanied by cyclical changes in the texture or staining properties of either nucleoplasm or cytoplasm; a microtubular (spindle) system is not formed. The plasma membrane (cytoplasmic membrane) is frequently complex in topology and forms vesicular, lamellar, or tubular intrusions into the cytoplasm; vacuoles and replicating cytoplasmic organelles independent of the plasma membrane system (chlorobium vesicles, gas vacuoles) are relatively rare and are enclosed by nonunit membranes. Respiratory and photosynthetic functions are associated with the plasma-membrane system in those members possessing these physiological attributes, although in the cyanobacteria there may be an independence of plasma and thylakoid membranes. Ribosomes of the 70S type (except for one groupthe Archaeawith slightly higher S values) are dispersed in the cytoplasm; an endoplasmic reticulum with attached ribosomes is not present. The cytoplasm is immobile; cytoplasmic streaming, pseudopodial movement, endocytosis, and exocytosis are not observed. Nutrients are acquired in molecular form. Enclosure of the cell by a rigid wall is common but not universal. The cell may be nonmotile or may exhibit swimming motility (mediated by flagella of bacterial type) or gliding motility on surfaces.

 In organismal terms, these ubiquitous inhabitants of moist environments are predominantly unicellular microorganisms, but filamentous, mycelial, or colonial forms also occur.
Differentiation is limited in scope (holdfast structures, resting cells, and modifications in cell shape). Mechanisms of gene transfer and recombination occur, but these processes never involv gametogenesis and zygote formation.

 Although procaryotic organisms can usually be readily differentiated from eucaryotic microorganisms, in some instances it may be difficult, especially with procaryotes that exhibit some attributes similar to those of microscopic eucaryotes. For instance,

The hyphae formed by actinomycetes might be confused with the hyphae formed by molds; A fascicle of bacterial flagella could give the misleading impression of being a single eucaryotic flagellum; The ability of spirochetes to twist and contort their shape is suggestive of the flexibility exhibited by certain protozoa; some eucaryotic cells are as small as bacteria, and some bacteria are as large as eucaryotic cells (see footnote to Table 1). ,

 The most reliable approach is probably the demonstration of the absence of a nuclear membrane in procaryotes.
Other procaryotic cell features range from those that are relatively easy to determine to the molecular characteristics that require sophisticated methods. Fluorescent- labeled gene probes that can easily distinguish between procaryotic and eucaryotic cells have been developed. Some characteristics that may help to differentiate between procaryotes and eucaryotes are listed in Table 1.

ARCHAEA VS. BACTERIA

The two fundamentally different groups (domains) that comprise the procaryotes are the Bacteria and the Archaea. Recent phylogenetic analyses of the Bacteria, Archaea, and Eucarya using conserved protein sequences have shown that the majority of trees support a closer relationship between the Archaea and Eucarya than between Archaea and Bacteria.

Although the possibility of interdomain horizontal gene transfer complicates this picture, the apparent Archaea Eucarya sisterhood raises interesting questions about the phylogenetic relationships between procaryotes and eucaryotes and the root of the universal tree of life. Table 2 provides some characteristics differentiating these two procaryotic groups. A general description of each group follows.

Bacteria

For practical purposes the Bacteria may be divided into three phenotypic subgroups: (1) those that are Gram-negative and have a cell wall, (2) those that are Gram-positive and have a cell wall, and (3) those that lack any cell wall. Gram-negative Bacteria that have a cell wall
These have a Gram-negative type of cell-wall profile consisting of an outer membrane and an inner, relatively thin peptidoglycan layer (which contains muramic acid and is present in all but a few organisms that have lost this portion of wall; see footnote to Table 1) as well as a variable complement of other components outside or between these layers. They usually stain Gram-negative, although the presence of a thick exopolysaccharide layer around the outer membrane may result in a Gram-positive staining reaction, as seen in the cyst-like forms of some Azospirillum species. Cell shapes may be spheres, ovals, straight or curved rods, helices, or filaments; some of these forms may be sheathed or capsulated. Reproduction is by binary fission, but some groups show budding, and a rare group (Subsection II of the cyanobacteria) shows multiple fission. Fruiting bodies and myxospores may be formed by the myxobacteria. Swimming motility, gliding motility, and nonmotility are commonly observed. Members may be phototrophic or nonphototrophic (both lithotrophic and heterotrophic) bacteria and include aerobic, anaerobic, facultatively anaerobic, and microaerophilic species;

Gram-positive Bacteria that have a cell wall

These Bacteria have a cell-wall profile of the Gram-positive type; there is no outer membrane and the peptidoglycan layer is relatively thick.
Some members of the group have teichoic acids and/or neutral polysaccharides as components of the wall. A few members of the group have cell walls that contain mycolic acids. Reaction with Grams stain is generally, but not always, positive; exceptions such as Butyrivibrio, which has an unusually thin wall and stains Gram-negative, may occur. Cells may be spheres, rods, or filaments; the rods and filaments may be nonbranching, but many show true branching.

Cellular reproduction is generally by binary fission; some produce spores as resting forms (endospores or spores on hyphae). The members of this division include simple asporogenous and sporogenous bacteria, as well as the actinomycetes and their relatives.
Gram-positive Bacteria are generally chemosynthetic heterotrophs and include aerobic, anaerobic, facultatively anaerobic, and microaerophilic species; somemembers are obligate intracellular parasites. Only one group, the heliobacteria, is photosynthetic and although these have a Gram-positive type of cell wall they nevertheless stain Gram- negative.

Bacteria lacking a cell wall

These Bacteria are commonl called the mycoplasmas. They do not synthesize the precursors of peptidoglycan and are insensitive to b-lactam antibiotics or other antibiotics that inhibit cell wall synthesis. They are enclosed by a unit membrane, the plasma membrane. The cells are highly pleomorphic and range in size from large deformable vesicles to very small (0.2 lm), filterable elements. Filamentous forms with branching projections are common. Reproduction may be by budding, fragmentation, and/or binary fission. Some groups show a degree of regularity of form due to the placing of internal structures. Usually, they are nonmotile, but some species show a form of gliding motility. No resting forms are known. Cells stain Gram-negative. Most require complex media for growth (high-osmotic-pressure surroundings) and tend to penetrate the surface of solid media forming characteristic fried egg colonies.

The organisms resemble the naked L-forms that can be generated from many species of bacteria (notably Grampositive Bacteria) but differ in that the mycoplasmas are unable to revert and make cell walls. Most species are further distinguished by requiring both cholesterol and long-chain fatty acids for growth; unesterified cholesterol is a unique compo
nent of the membranes of both sterol-requiring and nonrequiring species if present in the medium. The mol% G C content of rRNA is 4348 (lower than the 5054 mol% of walled Gramnegative and Gram-positive Bacteria); the mol% G C content of the DNA is also relatively low, 2346, and the genome size of the mycoplasmas at 0.51.0 109 Da is less than that of other procaryotes. The mycoplasmas may be saprophytic, parasitic, or pathogenic, and the pathogens cause diseases of animals, plants, and tissue cultures.