The cell envelope comprises the inner cell membrane and the cell wall of a bacterium. In gram-negative bacteria an outer membrane is also included. This envelope is not present in the Mollicutes where the cell wall is absent.
Bacterial cell envelopes fall into two major categories: a gram-positive type which stains purple during gram staining and a gram-negative type which stains pink during gram staining. Either type may have an enclosing capsule of polysaccharides for extra protection. As a group these are known as polysaccharide encapsulated bacteria.
Function
As in other organisms, the bacterial cell wall provides structural integrity to the cell. In prokaryotes, the primary function of the cell wall is to protect the cell from internal turgor pressure caused by the much higher concentrations of proteins and other molecules inside the cell compared to its external environment. The bacterial cell wall differs from that of all other organisms by the presence of peptidoglycan (poly-N-acetylglucosamine and N-acetylmuramic acid), which is located immediately outside of the cytoplasmic membrane. Peptidoglycan is responsible for the rigidity of the bacterial cell wall and for the determination of cell shape. It is relatively porous and is not considered to be a permeability barrier for small substrates. While all bacterial cell walls (with a few exceptions e.g. intracellular parasites such as Mycoplasma) contain peptidoglycan, not all cell walls have the same overall structures. This is notably expressed through the classification into gram positive and gram negative bacteria.
Types of bacterial cell envelopes
The gram-positive cell wall
The gram-positive cell wall is characterized by the presence of a very thick peptidoglycan layer, which is responsible for the retention of the crystal violet dyes during the Gram staining procedure. It is found exclusively in organisms belonging to the Actinomycetota (or high %G+C gram-positive organisms) and the Bacillota (or low %G+C gram-positive organisms). Bacteria within the Deinococcota group may also exhibit gram-positive staining behavior but contain some cell wall structures typical of gram-negative organisms. Imbedded in the gram-positive cell wall are polyalcohols called teichoic acids, some of which are lipid-linked to form lipoteichoic acids. Because lipoteichoic acids are covalently linked to lipids within the cytoplasmic membrane they are responsible for linking the peptidoglycan to the cytoplasmic membrane. Teichoic acids give the gram-positive cell wall an overall negative charge due to the presence of phosphodiester bonds between teichoic acid monomers.
Outside the cell wall, many Gram-positive bacteria have an S-layer of "tiled" proteins. The S-layer assists attachment and biofilm formation. Outside the S-layer, there is often a capsule of polysaccharides. The capsule helps the bacterium evade host phagocytosis. In laboratory culture, the S-layer and capsule are often lost by reductive evolution (the loss of a trait in absence of positive selection).
The gram-negative cell wall
The gram-negative cell wall contains a thinner peptidoglycan layer adjacent to the cytoplasmic membrane than the gram-positive wall, which is responsible for the cell wall's inability to retain the crystal violet stain upon decolourisation with ethanol during Gram staining. In addition to the peptidoglycan layer the gram-negative cell wall also contains an additional outer membrane composed by phospholipids and lipopolysaccharides which face into the external environment. The highly charged nature of lipopolysaccharides confer an overall negative charge to the gram -negative cell wall. The chemical structure of the outer membrane lipopolysaccharides is often unique to specific bacterial strains (i.e. sub-species) and is responsible for many of the antigenic properties of these strains.