
[New Latin E(scherichia) colī, species name : after Theodor Escherich (1857-1911), German physician + Latin colī, genitive of colon, colon; see colon2.]
For more information on E. coli, visit Britannica.com.
A group of bacteria including both harmless ones that inhabit human intestines and some types that can cause food poisoning.
The primary bacterial indicator used for assessment of microbial contamination of water consists of the coliform group. Coliform bacteria are universally present in high numbers in the feces of warm-blooded animals, including humans, and can be detected even after considerable dilution.
Escherichia coli (E. coli), is one of the most common coliform bacteria types. Detection of E. coli is definite evidence of fecal pollution. E. coli are facultatively anaerobic gram-negative rods that live in the intestinal tracts of animals. They can grow in the presence or the absence of oxygen. Under anaerobic conditions, E. coli grow by fermentation, producing mixed acids and gases as end products. They can also grow by anaerobic respiration, utilizing NO3, NO2, or fumarate. This versatility is what gives E. coli its ability to adapt to its intestinal (anaerobic) and its extraintestinal (aerobic or anaerobic) habitats.
As a pathogen, E. coli is best known for its ability to cause intestinal diseases. Five classes of E. coli can result in diarrheal diseases, but three specific pathogenic strains—enterotoxigenic, enteropathogenic, and enteroinvasive—cause problems when present in the water supply. All three of these types can cause acute diarrhea. An outbreak of E. coli-induced diarrhea can have a fatality rate as high as 40 percent in newborn children.
Enterotoxigenic E. coli (ETEC) are an important cause of diarrhea in infants (e.g., in nurseries and institutions), and in travelers to areas with poor sanitation. ETEC are acquired by ingestion of contaminated food and water. Adults in endemic areas develop immunity. In developing countries, children under the age of three experience multiple ETEC infections. The primary symptom of ETEC infection is diarrhea without fever.
Enteroinvasive E. coli (EIEC) penetrate and multiply within epithelial cells of the colon and cause widespread cell destruction. EIEC are very similar to Shigella in their pathogenic mechanisms and the type of clinical symptoms they cause—diarrhea with fever. EIEC infections are endemic in developing countries and are the cause of 1 to 5 percent of diarrheal episodes among people seeking treatment.
Enteropathogenic E. coli (EPEC) are an important cause of traveler's diarrhea in Mexico and in North America. This class of E. coli produces watery diarrhea similar to that of ETEC, probably due to the bacterial invasion of host cells and modification of cellular signals. Diarrheal episodes among children caused by EPEC in endemic populations are normally limited to children under the age of one. In this age group, EPEC causes watery diarrhea with mucus, fever, and dehydration. EPEC is no longer an important cause of infant diarrhea in North America and Europe, but is still a major cause in many developing countries in South America, southern Africa, and Asia.
Escherica coli 0157:H7 is classified by the Centers for Disease Control and Prevention as the cause of one of the emerging infections diseases. E. coli 0157:H7 is one of the more virulent of the many strains of E.coli found in the environment. (The CDC reports that 20,000 cases of 0157:H7 infection may occur annually.) E. coli 0157:H7 is found in the intestinal tract and feces of animals and humans. Infection often causes severe, bloody diarrhea and abdominal cramps. In children, the elderly, and immune-compromised individuals, the infection can lead to kidney failure and possible death. Undercooked ground beef (due to its handling and preparation) represents one of the greatest risks of E. coli 0157:H7 infections.
Bibliography
Todar, K. "Bacteriology 330 Lecture Topics: Pathogenic E.Coli." Available at http://www.md.huji.ac.il/microbiology/bact330/lectureecoli.html.
Wallace, R. (1998). Maxey-Rosenau-Last Public Health and Preventive Medicine, 14th edition. Stamford, CT: Appleton and Lange.
— MARK G. ROBSON
Escherichia coli (E. coli) is a very common bacterium that normally inhabits the digestive tract of animals, including humans. It is widespread in the natural world and can also be found in soil and water. It is a member of the bacterial family Enterobacteriaciae, which also includes the bacteria Shigella, Salmonella, and Yersinia, among others. Some of these organisms, including E. coli, can cause serious diseases under certain conditions.
Attributes of E. Coli
E. coli is important to human health because it is a source of vitamins B12 and K, which it manufactures from undigested food in the large intestine. Unlike many other intestinal bacteria, E. coli can survive and grow in the presence of oxygen (although it can also grow without oxygen), which makes it a useful experimental model organism in the laboratory.
Even though E. coli is a single species of bacteria, many different varieties (called strains) of the species exist. Each has different characteristics, and while some are safe model organisms, others can cause potentially deadly disease. This is the case with E. coli 0157:H7, which is considered a dangerous pathogen which can infect humans. This strain is significantly different from the commonly used laboratory strains, which do not cause disease.
Importance in Laboratory Studies
E. coli is the most well-understood bacterium in the world, and is an extremely important model organism in many fields of research, particularly molecular biology, genetics, and biochemistry. It is easy to grow under laboratory conditions, and research strains are very safe to work with. As with many bacteria, E. coli grows quickly, which allows many generations to be studied in a short time. In fact, under ideal conditions, E. coli cells can double in number after only 20 minutes.
Furthermore, a very large number of E. coli bacteria can be grown in a small space—many millions in a drop of broth, for example. These are important characteristics in genetic experiments, which often involve selecting a single bacterial cell from among millions of candidates, then allowing it to reproduce into high numbers again to perform additional experiments.
Many vital techniques, such as molecular cloning and overexpression of cloned genes, were initially developed in E. coli and are still simpler and more effective in the bacterium. Crucial experiments that illuminated the details of fundamental biological processes such as DNA replication, transcription, and translation were performed for the first time or with greatest success in E. coli. The bacterium is still a primary resource in many modern laboratories. Even research efforts that focus on other organisms, including humans or crop plants, often use E. coli extensively as a tool to facilitate cloning and DNA sequencing.
Discoveries Made in E. Coli
Some of the discoveries made in E. coli have provided an invaluable framework for understanding biological processes in more complex organisms. As mentioned above, many fundamental processes that are shared by all living things are most easily studied in this simple bacterial model. Furthermore,E. coli has served as a model for understanding the biology of other bacteria.
The ways in which E. coli interacts with the human body are in many cases very similar to the ways that other disease-causing organisms act. Therefore, this model organism has been important in the study of human health, and has allowed researchers to ask questions about bacteria in general (for example, how antibiotics stop infections, or how the immune system fights off disease).
Genome Sequenced Early
Sequencing of the E. coli K-12 strain genome (a popular model strain) was completed in 1997; subsequently, at least two collections of the pathogenic 0157:H7 strain have been completely sequenced. The bacterium has a genome of approximately 4.3 million base pairs of DNA, and carries about 4,400 genes. Interestingly, only about 50 percent of the predicted genes have been described and characterized, a surprisingly low percentage for such a well-understood organism. For this and other reasons, E. coli remains one of the most significant model organisms used today.
Bibliography
Madigan, Michael T., John M. Martinko, and Jack Parker. Brock Biology of Micro-organisms, 9th ed. Upper Saddle River, NJ: Prentice Hall, 2000.
—Daniel J. Tomso
E. Coli Food Poisoning
In 1982 a particularly toxic strain of E. coli, E. coli 0157:H7, was identified; it produces a toxin (Shiga toxin) that damages cells that line the intestines. The same strain was responsible for a 1993 outbreak of food poisoning in Washington state, which sickened 500 people and killed three, and a series of outbreaks in 1996 in Japan, which sickened some 10,000 and killed 12. A rarer and more virulent Shiga-toxin-producing strain, E. coli O104:H4, was the cause of the 2011 outbreak centered on N Germany that sickened more than 4,000 people from more than a dozen countries and killed 50.
Food-poisoning outbreaks due to Shiga-toxin-producing E. coli (STEC) are typically the result of transmission via raw or undercooked ground meat (thought to become contaminated during slaughter or processing) or contaminated salad ingredients. The strains can potentially contaminate any food. STEC infections can also occur through other means, such as contact with infected persons or cattle, or consumption of or contact with contaminated water.
Symptoms, which begin 1 to 8 days after infection and last for about a week, include bloody diarrhea, abdominal pain, vomiting, and in some cases, fever. The most serious complication is a hemolytic-uremic syndrome (HUS) that can lead to kidney failure and death, especially in children. The 2011 infection, for example, led to HUS in some 900 people. There is no treatment other than supportive care. Practical preventive measures include thorough cooking of meat and careful hygiene around infected individuals.
A rapid rise in the number of cases of illness caused by STEC strains has prompted calls for a reevaluation of food inspection techniques in the United States. Irradiation of meat and some greens is now approved by the FDA as a means to destroy such bacteria.
A species of coliform bacteria normally present in the intestines and common in water, milk, and soil.
| Escherichia coli | |
|---|---|
| Scientific classification | |
| Domain: | Bacteria |
| Kingdom: | Eubacteria |
| Phylum: | Proteobacteria |
| Class: | Gammaproteobacteria |
| Order: | Enterobacteriales |
| Family: | Enterobacteriaceae |
| Genus: | Escherichia |
| Species: | E. coli |
| Binomial name | |
| Escherichia coli (Migula 1895) Castellani and Chalmers 1919 |
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| Synonyms | |
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Bacillus coli communis Escherich 1885 |
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Escherichia coli (
/ˌɛʃɨˈrɪkiə ˈkoʊlaɪ/;[1] commonly abbreviated E. coli) is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms (endotherms). Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls due to food contamination.[2][3] The harmless strains are part of the normal flora of the gut, and can benefit their hosts by producing vitamin K2,[4] and by preventing the establishment of pathogenic bacteria within the intestine.[5][6]
E. coli and related bacteria constitute about 0.1% of gut flora,[7] and fecal-oral transmission is the major route through which pathogenic strains of the bacterium cause disease. Cells are able to survive outside the body for a limited amount of time, which makes them ideal indicator organisms to test environmental samples for fecal contamination.[8][9] There is, however, a growing body of research that has examined environmentally persistent E. coli which can survive for extended periods of time outside of the host.[10] The bacterium can also be grown easily and inexpensively in a laboratory setting, and has been intensively investigated for over 60 years. E. coli is the most widely studied prokaryotic model organism,[citation needed] and an important species in the fields of biotechnology and microbiology, where it has served as the host organism for the majority of work with recombinant DNA.
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Contents
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The genera Escherichia and Salmonella diverged around 102 million years ago (credibility interval: 57–176 mya), which coincides with the divergence of their hosts: the former being found in mammals and the latter in birds and reptiles.[11] This was followed by a split of the escherichian ancestor into five species (E. albertii, E. coli, E. fergusonii, E. hermannii and E. vulneris. The last E. coli ancestor split between 20 and 30 mya.[12]
In 1885, Theodor Escherich, a German pediatrician, first discovered this species in the feces of healthy individuals and called it Bacterium coli commune due to the fact it is found in the colon and early classifications of Prokaryotes placed these in a handful of genera based on their shape and motility (at that time Ernst Haeckel's classification of Bacteria in the kingdom Monera was in place[13]).[14] Bacterium coli was the type species of the now invalid genus Bacterium when it was revealed that the former type species ("Bacterium triloculare") was missing.[15] Following a revision of Bacteria it was reclassified as Bacillus coli by Migula in 1895[16] and later reclassified in the newly created genus Escherichia, named after its original discoverer.[17]
The genus belongs in a group of bacteria informally known as "coliforms", and is a member of the Enterobacteriaceae family ("the enterics") of the Gammaproteobacteria.[18]
In May 2011, one E. coli strain, Escherichia coli O104:H4, has been the subject of a bacterial outbreak that began in Germany. Certain strains of E. coli are a major cause of foodborne illness. The outbreak started when several people in Germany were infected with enterohemorrhagic E. coli (EHEC) bacteria, leading to hemolytic-uremic syndrome (HUS), a medical emergency that requires urgent treatment. The outbreak did not only concern Germany, but 11 other countries, including regions in North America. [19] On 30 June 2011 the German Bundesinstitut für Risikobewertung (BfR) (Federal Institute for Risk Assessment, a federal, fully legal entity under public law of the Federal Republic of Germany, an institute within the German Federal Ministry of Food, Agriculture and Consumer Protection) announced that seeds of fenugreek from Egypt were likely the cause of the EHEC outbreak.[20]
E. coli is Gram-negative, facultative anaerobic and non-sporulating. Cells are typically rod-shaped, and are about 2.0 microns (μm) long and 0.5 μm in diameter, with a cell volume of 0.6 – 0.7 (μm)3.[21][22] It can live on a wide variety of substrates. E. coli uses mixed-acid fermentation in anaerobic conditions, producing lactate, succinate, ethanol, acetate and carbon dioxide. Since many pathways in mixed-acid fermentation produce hydrogen gas, these pathways require the levels of hydrogen to be low, as is the case when E. coli lives together with hydrogen-consuming organisms, such as methanogens or sulphate-reducing bacteria.[23]
Optimal growth of E. coli occurs at 37°C (98.6°F) but some laboratory strains can multiply at temperatures of up to 49°C (120.2°F).[24] Growth can be driven by aerobic or anaerobic respiration, using a large variety of redox pairs, including the oxidation of pyruvic acid, formic acid, hydrogen and amino acids, and the reduction of substrates such as oxygen, nitrate, fumarate, dimethyl sulfoxide and trimethylamine N-oxide.[25]
Strains that possess flagella are motile. The flagella have a peritrichous arrangement.[26]
E. coli and related bacteria possess the ability to transfer DNA via bacterial conjugation, transduction or transformation, which allows genetic material to spread horizontally through an existing population. This process led to the spread of the gene encoding shiga toxin from Shigella to E. coli O157:H7, carried by a bacteriophage.[27]
Escherichia coli encompasses an enormous population of bacteria that exhibit a very high degree of both genetic and phenotypic diversity. Genome sequencing of a large number of isolates of E. coli and related bacteria shows that a taxonomic reclassification would be desirable. However, this has not been done, largely due to its medical importance[28] and Escherichia coli remains one of the most diverse bacterial species: only 20% of the genome is common to all strains.[29] In fact, from the evolutionary point of view, the members of genus Shigella (dysenteriae, flexneri, boydii, sonnei) should be classified as E. coli strains, a phenomenon termed taxa in disguise.[30] Similarly, other strains of E. coli (e.g. the K-12 strain commonly used in recombinant DNA work) are sufficiently different that they would merit reclassification.
A strain is a sub-group within the species that has unique characteristics that distinguish it from other strains. These differences are often detectable only at the molecular level; however, they may result in changes to the physiology or lifecycle of the bacterium. For example, a strain may gain pathogenic capacity, the ability to use a unique carbon source, the ability to take upon a particular ecological niche or the ability to resist antimicrobial agents. Different strains of E. coli are often host-specific, making it possible to determine the source of fecal contamination in environmental samples.[8][9] For example, knowing which E. coli strains are present in a water sample allows researchers to make assumptions about whether the contamination originated from a human, another mammal or a bird.
A common subdivision system of E. coli, but not based on evolutionary relatedness, is by serotype, which is based on major surface antigens (O antigen: part of lipopolysaccharide layer; H: flagellin; K antigen: capsule), e.g. O157:H7)[31] (NB: K-12, the common laboratory strain is not a serotype.)
Like all lifeforms, new strains of E. coli evolve through the natural biological processes of mutation, gene duplication and horizontal gene transfer, in particular 18% of the genome of the laboratory strain MG1655 was horizontally acquired since the divergence from Salmonella.[32] In microbiology, all strains of E. coli derive from E. coli K-12 or E. coli B strains. Some strains develop traits that can be harmful to a host animal. These virulent strains typically cause a bout of diarrhea that is unpleasant in healthy adults and is often lethal to children in the developing world.[33] More virulent strains, such as O157:H7 cause serious illness or death in the elderly, the very young or the immunocompromised.[5][33]
E. coli is the type species of the genus (Escherichia) and in turn Escherichia is the type genus of the family Enterobacteriaceae, where it should be noted that the family name does not stem from the genus Enterobacter + "i" (sic.) + "aceae", but from "enterobacterium" + "aceae" (enterobacterium being not a genus, but an alternative trivial name to enteric bacterium).[18][34][35]
The original strain described by Escherich is believed to be lost, consequently a new type strain (neotype) was chosen as a representative: the neotype strain is ATCC 11775, also known as NCTC 9001,[36] which is pathogenic to chickens and has a O1:K1:H7 serotype.[37] However, in most studies either O157:H7 or K-12 MG1655 or K-12 W3110 are used as a representative E.coli.
Escherichia coli is a species. A large number of strains belonging to this species have been isolated and characterised. In addition to serotype (vide supra), they can be classified according to their phylogeny, i.e. the inferred evolutionary history, as shown below where the species is divided into six groups.[29][38][39]
The link between phylogenetic distance ("relatedness") and pathology is small, e.g. the O157:H7 serotype strains, which form a clade ("an exclusive group") — group E below — are all enterohaerogic strains (EHEC), but not all EHEC strains are closely related. In fact, four different species of Shigella are nested among E. coli strains (vide supra), while Escherichia albertii and Escherichia fergusonii are outside of this group. All commonly used research strains of E. coli' belong to group A and are derived mainly from Clifton's K-12 strain (λ⁺ F⁺; O16) and to a lesser degree from d'Herelle's "Bacillus coli" strain (B strain)(O7).
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The first complete DNA sequence of an E. coli genome (laboratory strain K-12 derivative MG1655) was published in 1997. It was found to be a circular DNA molecule 4.6 million base pairs in length, containing 4288 annotated protein-coding genes (organized into 2584 operons), seven ribosomal RNA (rRNA) operons, and 86 transfer RNA (tRNA) genes. Despite having been the subject of intensive genetic analysis for approximately 40 years, a large number of these genes were previously unknown. The coding density was found to be very high, with a mean distance between genes of only 118 base pairs. The genome was observed to contain a significant number of transposable genetic elements, repeat elements, cryptic prophages, and bacteriophage remnants.[40]
Today, over 60 complete genomic sequences of Escherichia and Shigella species are available. Comparison of these sequences shows a remarkable amount of diversity; only about 20% of each genome represents sequences that are present in every one of the isolates, while approximately 80% of each genome can vary among isolates.[29] Each individual genome contains between 4,000 and 5,500 genes, but the total number of different genes among all of the sequenced E. coli strains (the pan-genome) exceeds 16,000. This very large variety of component genes has been interpreted to mean that two-thirds of the E. coli pan-genome originated in other species and arrived through the process of horizontal gene transfer.[41]
E. coli normally colonizes an infant's gastrointestinal tract within 40 hours of birth, arriving with food or water or with the individuals handling the child. In the bowel, it adheres to the mucus of the large intestine. It is the primary facultative anaerobe of the human gastrointestinal tract.[42] (Facultative anaerobes are organisms that can grow in either the presence or absence of oxygen.) As long as these bacteria do not acquire genetic elements encoding for virulence factors, they remain benign commensals.[43]
According to its manufacturer, nonpathogenic Escherichia coli strain Nissle 1917 also known as Mutaflor is used as a probiotic agent in medicine, mainly for the treatment of various gastroenterological diseases,[44] including inflammatory bowel disease.[45]
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Virulent strains of E. coli can cause gastroenteritis, urinary tract infections, and neonatal meningitis. In rarer cases, virulent strains are also responsible for hemolytic-uremic syndrome, peritonitis, mastitis, septicemia and Gram-negative pneumonia.[42]
Because of its long history of laboratory culture and ease of manipulation, E. coli also plays an important role in modern biological engineering and industrial microbiology.[46] The work of Stanley Norman Cohen and Herbert Boyer in E. coli, using plasmids and restriction enzymes to create recombinant DNA, became a foundation of biotechnology.[47]
E. coli is a very versatile host for the production of heterologous proteins,[48] and various protein expression systems have been developed which allow the production of recombinant proteins in E. coli. Researchers can introduce genes into the microbes using plasmids which permit high level expression of protein, and such protein may be mass produced in industrial fermentation processes. One of the first useful applications of recombinant DNA technology was the manipulation of E. coli to produce human insulin.[49] Many proteins previously thought difficult or impossible to be expressed in E. coli in folded form have also been successfully expressed in E. coli. For example, proteins with multiple disulphide bonds may be produced in the periplasmic space or in the cytoplasm of mutants rendered sufficiently oxidizing to allow disulphide-bonds to form,[50] while proteins requiring post-translational modification such as glycosylation for stability or function have been expressed using the N-linked glycosylation system of Campylobacter jejuni engineered into E. coli.[51][52][53]
Modified E. coli cells have been used in vaccine development, bioremediation, and production of immobilised enzymes.[48]
E. coli is frequently used as a model organism in microbiology studies. Cultivated strains (e.g. E. coli K12) are well-adapted to the laboratory environment, and, unlike wild type strains, have lost their ability to thrive in the intestine. Many lab strains lose their ability to form biofilms.[54][55] These features protect wild type strains from antibodies and other chemical attacks, but require a large expenditure of energy and material resources.
In 1946, Joshua Lederberg and Edward Tatum first described the phenomenon known as bacterial conjugation using E. coli as a model bacterium,[56] and it remains the primary model to study conjugation.[citation needed] E. coli was an integral part of the first experiments to understand phage genetics,[57] and early researchers, such as Seymour Benzer, used E. coli and phage T4 to understand the topography of gene structure.[58] Prior to Benzer's research, it was not known whether the gene was a linear structure, or if it had a branching pattern.
E. coli was one of the first organisms to have its genome sequenced; the complete genome of E. coli K12 was published by Science in 1997.[59]
The long-term evolution experiments using E. coli, begun by Richard Lenski in 1988, have allowed direct observation of major evolutionary shifts in the laboratory.[60] In this experiment, one population of E. coli unexpectedly evolved the ability to aerobically metabolize citrate, which is extremely rare in E. coli. As the inability to grow aerobically is normally used as a diagnostic criterion with which to differentiate E. coli from other, closely related bacteria, such as Salmonella, this innovation may mark a speciation event observed in the lab.
By evaluating the possible combination of nanotechnologies with landscape ecology, complex habitat landscapes can be generated with details at the nanoscale.[61] On such synthetic ecosystems, evolutionary experiments with E. coli have been performed to study the spatial biophysics of adaptation in an island biogeography on-chip.
Studies are also being performed into programming E. coli to potentially solve complicated mathematics problems, such as the Hamiltonian path problem.[62]
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