Enterobacteria Virus RB18
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Enterobacteriaceae is a large family of Gram-negative bacteria. It was first proposed by Rahn in 1936, and now includes over 30 genera and more than 100 species. Its classification above the level of family is still a subject of debate, but one classification places it in the order Enterobacterales of the class
Gammaproteobacteria Gammaproteobacteria is a class of bacteria in the phylum Pseudomonadota (synonym Proteobacteria). It contains about 250 genera, which makes it the most genera-rich taxon of the Prokaryotes. Several medically, ecologically, and scientifically imp ...
in the phylum
Pseudomonadota Pseudomonadota (synonym Proteobacteria) is a major phylum of Gram-negative bacteria. The renaming of phyla in 2021 remains controversial among microbiologists, many of whom continue to use the earlier names of long standing in the literature. The ...
. In 2016, the description and members of this family were emended based on comparative genomic analyses by Adeolu et al. Enterobacteriaceae includes, along with many harmless
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
, many of the more familiar pathogens, such as ''
Salmonella ''Salmonella'' is a genus of rod-shaped (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' is the type species and is fur ...
'', '' Escherichia coli'', '' Klebsiella'', and '' Shigella''. Other disease-causing bacteria in this family include '' Enterobacter'' and ''
Citrobacter ''Citrobacter'' is a genus of Gram-negative coliform bacteria in the family Enterobacteriaceae. The species ''C. amalonaticus'', ''C. koseri'', and ''C. freundii'' can use citrate as a sole carbon source. ''Citrobacter'' species are differentia ...
''. Members of the Enterobacteriaceae can be trivially referred to as enterobacteria or "enteric bacteria",as several members live in the intestines of animals. In fact, the etymology of the family is enterobacterium with the suffix to designate a family (aceae)—not after the genus '' Enterobacter'' (which would be "Enterobacteraceae")—and the type genus is '' Escherichia''.


Morphology

Members of the Enterobacteriaceae are bacilli (rod-shaped), and are typically 1–5 μm in length. They typically appear as medium to large-sized grey colonies on blood agar, although some can express pigments. Most have many
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
used to move about, but a few genera are nonmotile. Most members of Enterobacteriaceae have peritrichous, type I fimbriae involved in the adhesion of the bacterial cells to their hosts. They are not
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
-forming.


Metabolism

Like other Pseudomonadota, Enterobactericeae have Gram-negative stains, and they are facultative anaerobes, fermenting sugars to produce lactic acid and various other end products. Most also reduce
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
to
nitrite The nitrite polyatomic ion, ion has the chemical formula . Nitrite (mostly sodium nitrite) is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name ...
, although exceptions exist. Unlike most similar bacteria, Enterobacteriaceae generally lack cytochrome c oxidase, there are exceptions. Catalase reactions vary among Enterobacteriaceae.


Ecology

Many members of this family are normal members of the
gut microbiota Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut m ...
in humans and other animals, while others are found in water or soil, or are parasites on a variety of different animals and plants.


Model organisms and medical relevance

'' Escherichia coli'' is one of the most important
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
s, and its genetics and biochemistry have been closely studied. Some enterobacteria are important pathogens, e.g. ''Salmonella'', or ''Shigella'' e.g. because they produce endotoxins. Endotoxins reside in the cell wall and are released when the cell dies and the cell wall disintegrates. Some members of the ''Enterobacteriaceae'' produce endotoxins that, when released into the bloodstream following cell lysis, cause a systemic inflammatory and vasodilatory response. The most severe form of this is known as endotoxic shock, which can be rapidly fatal.


Historical systematics and taxonomy

''Enterobacteriaceae'' was originally the sole family under the order 'Enterobacteriales'. The family contained a large array of biochemically distinct species with different ecological niches, which made biochemical descriptions difficult. The original classification of species to this family and order was largely based on 16S rRNA genome sequence analyses, which is known to have low discriminatory power and the results of which changes depends on the algorithm and organism information used. Despite this, the analyses still exhibited polyphyletic branching, indicating the presence of distinct subgroups within the family. In 2016, the order 'Enterobacteriales' was renamed to Enterobacterales, and divided into 7 new families, including the emended ''Enterobacteriaceae'' family. This emendation restricted the family to include only those genera directly related to the type genus, which included most of the enteric species under the order. This classification was proposed based on the construction of several robust phylogenetic trees using conserved genome sequences, 16S rRNA sequences and multilocus sequence analyses. Molecular markers, specifically conserved signature indels, specific to this family were identified as evidence supporting the division independent of phylogenetic trees. In 2017, a subsequent study using comparative phylogenomic analyses identified the presence of 6 subfamily level clades within the family ''Enterobacteriaceae'', namely the “Escherichia clade”, “Klebsiella clade”, “Enterobacter clade”, “Kosakonia clade”, “Cronobacter clade”, “Cedecea clade” and a “Enterobacteriaceae incertae sedis clade” containing species whose taxonomic placement within the family is unclear. However, this division was not officially proposed as the subfamily rank is generally not used.


Molecular signatures

Analyses of genome sequences from ''Enterobacteriaceae'' species identified 21 conserved signature indels (CSIs) that are uniquely present in this family in the proteins NADH:ubiquinone oxidoreductase (subunit M), twitching motility protein PilT, 2,3-dihydroxybenzoate-AMP ligase, ATP/GTP-binding protein, multifunctional fatty acid oxidation complex (subunit alpha),
S-formylglutathione hydrolase The enzyme ''S''-formylglutathione hydrolase (EC 3.1.2.12) catalyzes the reaction :S-formylglutathione + H2O \rightleftharpoons glutathione + formate This enzyme belongs to the family of hydrolases, specifically those acting on thioester bonds. ...
, aspartate-semialdehyde dehydrogenase,
epimerase Epimerases and racemases are isomerase enzymes that catalyze the inversion of stereochemistry in biological molecules. Racemases catalyze the stereochemical inversion around the asymmetric carbon atom in a substrate having only one center of asymme ...
, membrane protein, formate dehydrogenylase (subunit 7), glutathione S-transferase, major facilitator superfamily transporter,
phosphoglucosamine mutase In enzymology, a phosphoglucosamine mutase () is an enzyme that catalyzes the chemical reaction :alpha-D-glucosamine 1-phosphate \rightleftharpoons D-glucosamine 6-phosphate Hence, this enzyme has one substrate, alpha-D-glucosamine 1-phosphate, ...
, glycosyl hydrolase 1 family protein, 23S rrna racil(1939)-C(5)methyltransferase, co-chaperone HscB, N-acetylmuramoyl-L-alanine amidase, sulfate ABC transporter ATP-binding protein CysA, and LPS assembly protein LptD. These CSIs provide a molecular means of distinguishing ''Enterobacteriaceae'' from other families within the order ''Enterobacterales'' and other bacteria.


Genera


Validly published genera

The following genera have been validly published, thus they have "Standing in Nomenclature". The year the genus was proposed is listed in parentheses after the genus name. *''
Biostraticola ''Biostraticola'' is a Gram-negative, rod-shaped, non- endospore-forming, facultatively anaerobic and motile genus of bacteria within the family of Pectobacteriaceae The ''Pectobacteriaceae'' are a family of Gram-negative bacteria which largel ...
'' (2008) *''
Buttiauxella ''Buttiauxella'' is a Gram-negative, aerobic, facultative anaerobic and motile genus of bacteria within the family of Enterobacteriaceae Enterobacteriaceae is a large family (biology), family of Gram-negative bacteria. It was first propo ...
'' (1982) *''
Cedecea ''Cedecea'' is a genus of extremely rare bacteria of the family Enterobacteriaceae. The name of this genus was derived from CDC, the abbreviation for the Centers for Disease Control where the initial members of this genus were discovered. This ...
'' (1981) *''
Citrobacter ''Citrobacter'' is a genus of Gram-negative coliform bacteria in the family Enterobacteriaceae. The species ''C. amalonaticus'', ''C. koseri'', and ''C. freundii'' can use citrate as a sole carbon source. ''Citrobacter'' species are differentia ...
'' (1932) *''
Cronobacter ''Cronobacter'' is a genus of Gram-negative, facultatively anaerobic, oxidase-negative, catalase-positive, rod-shaped bacteria of the family Enterobacteriaceae. Several ''Cronobacter'' species are desiccation resistant and persistent in dry ...
'' (2008) *'' Enterobacillus'' (2015) *'' Enterobacter'' (1960) *'' Escherichia'' (1919) *'' Franconibacter'' (2014) *'' Gibbsiella'' (2011) *'' Izhakiella'' (2016) *'' Klebsiella'' (1885) *'' Kluyvera'' (1981) *'' Kosakonia'' (2013) *'' Leclercia'' (1987) *'' Lelliottia'' (2013) *'' Limnobaculum'' (2018) *''
Mangrovibacter ''Mangrovibacter'' is a genus in the order Enterobacterales. Members of the genus are Gram-stain-negative, facultatively anaerobic, nitrogen-fixing, and rod shaped. The name ''Mangrovibacter'' derives from:Neo-Latin noun ''mangrovum'', mangrove ...
'' (2010) *'' Metakosakonia'' (2017) *'' Phytobacter'' (2017) *''
Pluralibacter ''Pluralibacter'' is a genus of Gram negative bacteria from the family of Enterobacteriaceae Enterobacteriaceae is a large family (biology), family of Gram-negative bacteria. It was first proposed by Rahn in 1936, and now includes over 30 ...
'' (2013) *''
Pseudescherichia ''Pseudescherichia'' is a Gram-negative genus of non-spore-forming, facultatively anaerobic rod-shaped bacteria from the family Enterobacteriaceae. Based on conserved signature indels (CSIs) differentiating it from other members of this family, ...
'' (2017) *'' Pseudocitrobacter'' (2014) *''
Raoultella The genus ''Raoultella'' is composed of Gram-negative, Oxidase test, oxidase-negative, Aerobic organism, aerobic, motility, nonmotile, Capsule (microbiology), capsulated, Facultative anaerobic organism, facultatively anaerobic rods (formerly desi ...
'' (2001) *'' Rosenbergiella'' (2013) *'' Saccharobacter'' (1990) *''
Salmonella ''Salmonella'' is a genus of rod-shaped (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' is the type species and is fur ...
'' (1900) *'' Scandinavium'' (2020) *'' Shigella'' (1919) *'' Shimwellia'' (2010) *'' Siccibacter'' (2014) *'' Trabulsiella'' (1992) *'' Yokenella'' (1985)


'' Candidatus'' genera

* "''Candidatus'' Annandia" * "''Candidatus'' Arocatia" * "''Candidatus'' Aschnera" * "''Candidatus'' Benitsuchiphilus" * "''Candidatus'' Blochmannia" * "''Candidatus'' Curculioniphilus" * "''Candidatus'' Cuticobacterium" * "''Candidatus'' Doolittlea" * "''Candidatus'' Gillettellia" * "''Candidatus'' Gullanella" * "''Candidatus'' Hamiltonella" * "''Candidatus'' Hartigia" * "''Candidatus'' Hoaglandella" * "''Candidatus'' Ischnodemia" * "''Candidatus'' Ishikawaella" * "''Candidatus'' Kleidoceria" * "''Candidatus'' Kotejella" * "''Candidatus'' Macropleicola" * "''Candidatus'' Mikella" * "''Candidatus'' Moranella" * "''Candidatus'' Phlomobacter" * "''Candidatus'' Profftia" * "''Candidatus'' Purcelliella" * "''Candidatus'' Regiella" * "''Candidatus'' Riesia" * "''Candidatus'' Rohrkolberia" * "''Candidatus'' Rosenkranzia" * "''Candidatus'' Schneideria" * "''Candidatus'' Stammera" * "''Candidatus'' Stammerula" * "''Candidatus'' Tachikawaea" * "''Candidatus'' Westeberhardia"


Proposed genera

The following genera have been effectively, but not validly, published, thus they do not have "Standing in Nomenclature". The year the genus was proposed is listed in parentheses after the genus name. *''Aquamonas'' (2009) *''Atlantibacter'' (2016) *'' Superficieibacter'' (2018)


Identification

To identify different
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
of Enterobacteriaceae, a microbiologist may run a series of tests in the lab. These include: * Phenol red *
Tryptone Tryptone is the assortment of peptides formed by the digestion of casein by the protease trypsin. Tryptone is commonly used in microbiology to produce lysogeny broth (LB) for the growth of '' E. coli'' and other microorganisms. It provides a so ...
broth *
Phenylalanine Phenylalanine (symbol Phe or F) is an essential α-amino acid with the formula . It can be viewed as a benzyl group substituted for the methyl group of alanine, or a phenyl group in place of a terminal hydrogen of alanine. This essential amino a ...
agar for detection of production of
deaminase Deamination is the removal of an amino group from a molecule. Enzymes that catalyse this reaction are called deaminases. In the human body, deamination takes place primarily in the liver, however it can also occur in the kidney. In situations of ...
, which converts phenylalanine to
phenylpyruvic acid Phenylpyruvic acid is the organic compound with the formula C6H5CH2C(O)CO2H. It is a keto acid. Occurrence and properties The compound exists in equilibrium with its E- and Z-enol tautomers. It is a product from the oxidative deamination of phe ...
* Methyl red or Voges-Proskauer tests depend on the digestion of glucose. The methyl red tests for acid endproducts. The Voges Proskauer tests for the production of
acetylmethylcarbinol Acetoin, also known as 3-hydroxybutanone or acetyl methyl carbinol, is an organic compound with the formula CH3CH(OH)C(O)CH3. It is a colorless liquid with a pleasant, buttery odor. It is chiral. The form produced by bacteria is (''R'')-acetoin. ...
. *
Catalase Catalase is a common enzyme found in nearly all living organisms exposed to oxygen (such as bacteria, plants, and animals) which catalyzes the decomposition of hydrogen peroxide to water and oxygen. It is a very important enzyme in protecting t ...
test on
nutrient agar Nutrient agar is a general purpose liquid medium supporting growth of a wide range of non-fastidious organisms. It typically contains ( mass/volume): * 0.5% peptone - this provides organic nitrogen * 0.3% beef extract/yeast extract - the wate ...
tests for the production of enzyme catalase, which splits hydrogen peroxide and releases oxygen gas. *
Oxidase test The oxidase test is used to determine if an organism possesses the cytochrome c oxidase enzyme. The test is used as an aid for the differentiation of ''Neisseria'', ''Moraxella'', '' Campylobacter'' and ''Pasteurella'' species (oxidase positive). I ...
on
nutrient agar Nutrient agar is a general purpose liquid medium supporting growth of a wide range of non-fastidious organisms. It typically contains ( mass/volume): * 0.5% peptone - this provides organic nitrogen * 0.3% beef extract/yeast extract - the wate ...
tests for the production of the enzyme
oxidase In biochemistry, an oxidase is an enzyme that catalyzes oxidation-reduction reactions, especially one involving dioxygen (O2) as the electron acceptor. In reactions involving donation of a hydrogen atom, oxygen is reduced to water (H2O) or hydro ...
, which reacts with an aromatic amine to produce a purple color. * Nutrient gelatin tests to detect activity of the enzyme gelatinase. In a clinical setting, three species make up 80 to 95% of all isolates identified. These are ''Escherichia coli'', ''Klebsiella pneumoniae'', and ''Proteus mirabilis''. However, ''Proteus mirabilis'' is now considered a part of the
Morganellaceae The Morganellaceae are a family of Gram-negative bacteria that include some important human pathogens formerly classified as Enterobacteriaceae. This family is a member of the order Enterobacterales in the class Gammaproteobacteria of the phylum ...
, a sister clade within the Enterobacterales.


Antibiotic resistance

Several Enterobacteriaceae strains have been isolated which are resistant to antibiotics including carbapenems, which are often claimed as "the last line of antibiotic defense" against resistant organisms. For instance, some '' Klebsiella pneumoniae'' strains are carbapenem resistant. Various carbapenemases genes (blaOXA-48, blaKPC and blaNDM-1, blaVIM and blaIMP) have been identified in carbapenem resistant Enterobacteriaceae including ''Escherichia coli'' and ''Klebsiella pneumoniae''.


References


External links

* *
Enterobacteriaceae
genomes and related information a
PATRIC
a Bioinformatics Resource Center funded b
NIAID
* Evaluation of new computer-enhanced identification program for microorganisms: adaptation of BioBASE for identification of members of the family Enterobacteriacea

* Brown, A.E. (2009). Benson's microbiological applications: laboratory manual in general microbiology. New York: McGraw- Hill. {{Taxonbar, from=Q380136 Enterobacteriaceae, Bacteria families Gram-negative bacteria