Gram-negative bacteria

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Microscopic image of gram-negative Pseudomonas aeruginosa bacteria (pink-red rods)

Gram-negative bacteria are

cell wall sandwiched between an inner (cytoplasmic) membrane and an outer membrane.[citation needed] These bacteria are found in all environments that support life on Earth
.

Within this category, notable species include the

Several

classes of antibiotics have been developed to target gram-negative bacteria, including aminopenicillins, ureidopenicillins, cephalosporins, beta-lactam-betalactamase inhibitor combinations (such as piperacillin-tazobactam), folate antagonists, quinolones, and carbapenems. Many of these antibiotics also cover gram-positive bacteria. The antibiotics that specifically target gram-negative organisms include aminoglycosides, monobactams (such as aztreonam), and ciprofloxacin
.

Characteristics

Gram-negative cell wall structure
Gram-positive and -negative bacteria are differentiated chiefly by their cell wall
structure

Conventional gram-negative (LPS-diderm) bacteria display the following characteristics:[citation needed]

Classification

Along with cell shape, Gram staining is a rapid diagnostic tool and once was used to group species at the subdivision of Bacteria. Historically, the kingdom Monera was divided into four divisions based on Gram staining: Firmacutes (+), Gracillicutes (−), Mollicutes (0) and Mendocutes (var.).[3] Since 1987, the

Cavalier-Smith still treat them as a monophyletic taxon (though not a clade; his definition of monophyly requires a single common ancestor but does not require holophyly, the property that all descendants be encompassed by the taxon) and refer to the group as a subkingdom "Negibacteria".[5]

Taxonomy

Bacteria are traditionally classified based on their

monoderm bacteria, while gram-negative bacteria, having two membranes, are also known as diderm bacteria. It was traditionally thought that the groups represent lineages, i.e., the extra membrane only evolved once, such that gram-negative bacteria are more closely related to one another than to any gram-positive bacteria. While this is often true, the classification system breaks down in some cases, with lineage groupings not matching the staining result.[6][7][8][9] Thus, Gram staining cannot be reliably used to assess familial relationships of bacteria. Nevertheless, staining often gives reliable information about the composition of the cell membrane, distinguishing between the presence or absence of an outer lipid membrane.[6][10]

Of these two structurally distinct groups of

antibiotics and that the gram-negative bacteria are, in general, resistant to antibiotics, it has been proposed that the outer cell membrane in gram-negative bacteria (diderms) evolved as a protective mechanism against antibiotic selection pressure.[6][7][10][11] Some bacteria such as Deinococcus, which stain gram-positive due to the presence of a thick peptidoglycan layer, but also possess an outer cell membrane are suggested as intermediates in the transition between monoderm (gram-positive) and diderm (gram-negative) bacteria.[6][11] The diderm bacteria can also be further differentiated between simple diderms lacking lipopolysaccharide (LPS); the archetypical diderm bacteria, in which the outer cell membrane contains lipopolysaccharide; and the diderm bacteria, in which the outer cell membrane is made up of mycolic acid (e. g. Mycobacterium).[8][9][11][12]

The conventional LPS-diderm group of gram-negative bacteria (e.g.,

HSP60 (GroEL) protein. In addition, a number of bacterial taxa (including Negativicutes, Fusobacteriota, Synergistota, and Elusimicrobiota) that are either part of the phylum Bacillota (a monoderm group) or branches in its proximity are also found to possess a diderm cell structure.[9][11][12] They lack the GroEL signature.[11] The presence of this CSI in all sequenced species of conventional lipopolysaccharide-containing gram-negative bacterial phyla provides evidence that these phyla of bacteria form a monophyletic clade and that no loss of the outer membrane from any species from this group has occurred.[11]

Example species

The

]

Medically-relevant gram-negative

Medically relevant gram-negative

]

Gram-negative bacteria associated with

intensive-care units
.

Bacterial transformation

Transformation is one of three processes for horizontal gene transfer, in which exogenous genetic material passes from one bacterium to another, the other two being conjugation (transfer of genetic material between two bacterial cells in direct contact) and transduction (injection of foreign DNA by a bacteriophage virus into the host bacterium).[18][19] In transformation, the genetic material passes through the intervening medium, and uptake is completely dependent on the recipient bacterium.[18]

As of 2014 about 80 species of bacteria were known to be capable of transformation, about evenly divided between

gram-positive and gram-negative bacteria; the number might be an overestimate since several of the reports are supported by single papers.[18] Transformation has been studied in medically important gram-negative bacteria species such as Helicobacter pylori, Legionella pneumophila, Neisseria meningitidis, Neisseria gonorrhoeae, Haemophilus influenzae and Vibrio cholerae.[20] It has also been studied in gram-negative species found in soil such as Pseudomonas stutzeri, Acinetobacter baylyi, and gram-negative plant pathogens such as Ralstonia solanacearum and Xylella fastidiosa.[20]

Role in disease

MALDI-TOF
and/or multitarget assay.

One of the several unique characteristics of gram-negative bacteria is the structure of the

innate immune response, activating the immune system and producing cytokines (hormonal regulators). This leads to inflammation and can cause a toxic reaction, resulting in fever, an increased respiratory rate, and low blood pressure. That is why some infections with gram-negative bacteria can lead to life-threatening septic shock.[2]

The outer membrane protects the bacteria from several

beta-lactamase inhibitors to combat the presence of enzymes that can digest these drugs (known as beta-lactamases) in the peri-plasmic space. Other classes of drugs that have gram negative spectrum include cephalosporins, monobactams (aztreonam), aminoglycosides, quinolones, macrolides, chloramphenicol, folate antagonists, and carbapenems.[21]

Orthographic note

The adjectives gram-positive and gram-negative derive from the surname of

Gram staining § Orthographic note
.

See also

References

Notes

External links