Gram-negative bacteria are bacteria that, unlike gram-positive bacteria , do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is their cell envelope , which consists of a thin peptidoglycan cell wall sandwiched between an inner ( cytoplasmic ) membrane and an outer membrane . These bacteria are found in all environments that support life on Earth .
30-582: Pluralibacter is a genus of Gram negative bacteria from the family of Enterobacteriaceae . The genus consists of two species, P. gergoviae and P. pyrinus . Both species were originally classified in the genus Enterobacter but were reclassified into the novel genus Pluralibacter in 2013. Gram negative Within this category, notable species include the model organism Escherichia coli , along with various pathogenic bacteria , such as Pseudomonas aeruginosa , Chlamydia trachomatis , and Yersinia pestis . They pose significant challenges in
60-459: A monophyletic clade and that no loss of the outer membrane from any species from this group has occurred. The proteobacteria are a major superphylum of gram-negative bacteria, including E. coli , Salmonella , Shigella , and other Enterobacteriaceae , Pseudomonas , Moraxella , Helicobacter , Stenotrophomonas , Bdellovibrio , acetic acid bacteria , Legionella etc. Other notable groups of gram-negative bacteria include
90-427: 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". Bacteria are traditionally classified based on their Gram-staining response into the gram-positive and gram-negative bacteria. Having just one membrane,
120-657: A few conserved signature indel (CSI) in the 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. They lack the GroEL signature. 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
150-652: A multitude of species. Some of them cause primarily respiratory problems ( Klebsiella pneumoniae , Legionella pneumophila , Pseudomonas aeruginosa ), primarily urinary problems ( Escherichia coli , Proteus mirabilis , Enterobacter cloacae , Serratia marcescens ), and primarily gastrointestinal problems ( Helicobacter pylori , Salmonella enteritidis , Salmonella typhi ). Gram-negative bacteria associated with hospital-acquired infections include Acinetobacter baumannii , which cause bacteremia , secondary meningitis , and ventilator-associated pneumonia in hospital intensive-care units . Transformation
180-435: A number of different observations, including that the gram-positive bacteria are the most sensitive to 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 . Some bacteria such as Deinococcus , which stain gram-positive due to
210-787: A toxic reaction when the bacteria are lysed by immune cells. This reaction may lead to septic shock , resulting in low blood pressure , respiratory failure , reduced oxygen delivery , and lactic acidosis . 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 . Conventional gram-negative (LPS-diderm) bacteria display
240-519: 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 . The outer membrane protects the bacteria from several antibiotics , dyes , and detergents that would normally damage either the inner membrane or the cell wall (made of peptidoglycan ). The outer membrane provides these bacteria with resistance to lysozyme and penicillin . The periplasmic space (space between
270-403: 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). In transformation, the genetic material passes through
300-454: The CDC ), if any, governs the document being written. This is further explained at Gram staining § Orthographic note . Bacterial outer membrane The bacterial outer membrane is found in gram-negative bacteria . Gram-negative bacteria form two lipid bilayers in their cell envelopes - an inner membrane (IM) that encapsulates the cytoplasm , and an outer membrane (OM) that encapsulates
330-474: The cyanobacteria , spirochaetes , green sulfur , and green non-sulfur bacteria . Medically-relevant gram-negative diplococci include the four types that cause a sexually transmitted disease ( Neisseria gonorrhoeae ), a meningitis ( Neisseria meningitidis ), and respiratory symptoms ( Moraxella catarrhalis , A coccobacillus Haemophilus influenzae is another medically relevant coccal type. Medically relevant gram-negative bacilli include
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#1732855420031360-596: The periplasm . The composition of the outer membrane is distinct from that of the inner cytoplasmic cell membrane - among other things, the outer leaflet of the outer membrane of many gram-negative bacteria includes a complex lipopolysaccharide whose lipid portion acts as an endotoxin - and in some bacteria such as E. coli it is linked to the cell's peptidoglycan by Braun's lipoprotein . Porins can be found in this layer. Outer membrane proteins are membrane proteins with key roles associated with bacterial cell structure and morphology; cell membrane homeostasis;
390-632: The O-antigen transporter Wzx (RfbX) and the ATP binding cassette (ABC) transporter MsbA that flips the lipid A-core moiety from the inner leaflet to the outer leaflet of the inner membrane. O-antigen repeat units are then polymerised in the periplasm by the Wzy polymerase and ligated to the lipid A-core moiety by the WaaL ligase . The LPS transport machinery is composed of LptA, LptB, LptC, LptD, LptE. This supported by
420-420: The diderm bacteria in which the outer cell membrane is made up of mycolic acid (e. g. Mycobacterium ). The conventional LPS- diderm group of gram-negative bacteria (e.g., Pseudomonadota , Aquificota , Chlamydiota , Bacteroidota , Chlorobiota , " Cyanobacteria ", Fibrobacterota , Verrucomicrobiota , Planctomycetota , Spirochaetota , Acidobacteriota ; " Hydrobacteria ") are uniquely identified by
450-449: The fact that depletion of any one of these proteins blocks the LPS assembly pathway and results in very similar outer membrane biogenesis defects. Moreover, the location of at least one of these five proteins in every cellular compartment suggests a model for how the LPS assembly pathway is organised and ordered in space. LptC is required for the translocation of lipopolysaccharide (LPS) from
480-518: The following characteristics : 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.). Since 1987, the monophyly of the gram-negative bacteria has been disproven with molecular studies . However some authors, such as Cavalier-Smith still treat them as
510-487: The gram-positive bacteria are also known as 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
540-524: The individual components are transported from the site of synthesis to their final destination outside the inner membrane by crossing both hydrophilic and hydrophobic compartments. The machinery and the energy source that drive this process are not yet fully understood. The lipid A -core moiety and the O-antigen repeat units are synthesized at the cytoplasmic face of the inner membrane and are separately exported via two independent transport systems, namely,
570-411: The inner membrane to the outer membrane. LptE forms a complex with LptD, which is involved in the assembly of LPS in the outer leaflet of the outer membrane and is essential for envelope biogenesis. If lipid A , part of the lipopolysaccharide, enters the circulatory system it causes a toxic reaction by activating toll like receptor TLR 4 . Lipid A is very pathogenic and not immunogenic. However,
600-830: The intervening medium, and uptake is completely dependent on the recipient bacterium. 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. 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 . 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 . One of
630-440: The medical field due to their outer membrane, which acts as a protective barrier against numerous antibiotics (including penicillin ), detergents that would normally damage the inner cell membrane, and the antimicrobial enzyme lysozyme produced by animals as part of their innate immune system . Furthermore, the outer leaflet of this membrane contains a complex lipopolysaccharide (LPS) whose lipid A component can trigger
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#1732855420031660-402: The membrane called porins , and actively pumping efflux channels. The outer membranes of a bacterium can contain a huge number of proteins. In E. Coli for example there are around 500,000 in the membrane. Bacterial outer membrane proteins typically have a unique beta barrel structure that spans the membrane. The beta barrels fold to expose a hydrophobic surface before their insertion into
690-429: The outer membrane. Beta barrels vary in sequence and size that ranges from 8 to 36 beta strands. A subset of OMPs have a perisplasmic or an extracellular link to their beta barrel structure. An outer membrane protein is translocated across the inner membrane through ‘’Sec’’ machinery, and finally inserted to the outer membrane by the barrel assembly machinery complex. The biogenesis of the outer membrane requires that
720-509: 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 . The adjectives gram-positive and gram-negative derive from the surname of Hans Christian Gram , a Danish bacteriologist; as eponymous adjectives , their initial letter can be either capital G or lower-case g , depending on which style guide (e.g., that of
750-420: The polysaccharide component is very immunogenic, but not pathogenic, causing an aggressive response by the immune system. The sufferer will have a high temperature and respiration rate and a low blood pressure. This may lead to endotoxic shock , which may be fatal. The bacterial outer membrane is physiologically shed as the bounding membrane of outer membrane vesicles in cultures, as well as in animal tissues at
780-419: 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. 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
810-459: The several unique characteristics of gram-negative bacteria is the structure of the bacterial outer membrane . The outer leaflet of this membrane contains lipopolysaccharide (LPS), whose lipid A portion acts as an endotoxin . If gram-negative bacteria enter the circulatory system , LPS can trigger an innate immune response , activating the immune system and producing cytokines (hormonal regulators). This leads to inflammation and can cause
840-426: The staining result. 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 . Of these two structurally distinct groups of prokaryotic organisms, monoderm prokaryotes are thought to be ancestral. Based upon
870-403: The two cell membranes) also contains enzymes which break down or modify antibiotics. Drugs commonly used to treat gram negative infections include amino, carboxy and ureido penicillins ( ampicillin , amoxicillin , pipercillin , ticarcillin ). These drugs may be combined with beta-lactamase inhibitors to combat the presence of enzymes that can digest these drugs (known as beta-lactamases ) in
900-417: The uptake of nutrients; protection of the cell from toxins including antibiotics; and virulence factors including adhesins, exotoxins, and biofilm formation. There are a number of outer membrane proteins that are specifically virulence-related . Outer membrane proteins consist of two major classes of protein - transmembrane proteins and lipoproteins. The transmembrane proteins form channels or pores in
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