Vibrio cholerae

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Vibrio cholerae
Vibrio cholerae

Vibrio cholerae

Systematics
Department : Proteobacteria
Class : Gammaproteobacteria
Order : Vibrional
Family : Vibrionaceae
Genre : Vibrio
Type : Vibrio cholerae
Scientific name
Vibrio cholerae
Pacini 1854

Vibrio cholerae (formerly Vibrio comma ) is the causative agent of cholera . It is a gram-negative bacteria from the genus of vibrios . The cells are facultatively anaerobic , they can live with or without oxygen. The pathogen was in 1854 by Filippo Pacini as a curved, comma-shaped and highly portable bacterium described . In the same year, the Catalan Joaquim Balcells i Pascual described the pathogen. Robert Koch and Bernhard Fischer and Georg Gaffky cultivatedthe pathogenin pure culture from the intestines of deceased patients inIndia.

The species includes numerous strains of bacteria . The genome of the strain Vibrio cholerae O1 N16961 (also known as Biovar El Tor ) was completely sequenced in 2000 . It is distributed across two chromosomes , which is unusual for bacteria because most bacteria only have a single, circular bacterial chromosome. Not all bacterial strains are pathogenic ("disease-causing"). The bacterium acquires its pathogenicity through infection with bacteriophages ( viruses that specialize in bacteria).

features

Appearance

Light microscope image of Vibrio cholerae in a flagellum coloration according to Leifson (digitally colored afterwards)
Electron micrograph of Vibrio cholerae

As a typical representative of the genus Vibrio , Vibrio cholerae has the cell shape of a comma-shaped curved rod . In the Gram staining , it is gram-negative, i.e. it is colored red by the dyes used. This is caused by a thin layer of murein in the cell wall . He moves like other Vibrio - species with a single scourge continues. This sits at only one end of the bacterial cell, so that there is a monopolar monotriche flagellation . Forms of persistence such as endospores are not formed. Most strains do not have a capsule attached to the bacterial cell wall, with the exception of serogroup O139 , which has a capsule.

Growth and metabolism

The cells are facultatively anaerobic , so they can multiply even if there is no oxygen . They are catalase- positive and oxidase- positive, the latter serves as a distinguishing feature to representatives of the Enterobacteriaceae . The growth temperature in the natural habitat is 20–30 ° C, so Vibrio cholerae is one of the mesophilic (medium temperature preferring) bacteria. However, it can also tolerate slightly higher temperatures during growth. This is used if it is specifically cultivated as part of a microbiological examination. In most cases, a temperature of 35–42 ° C is used for cultivation . Since V. cholerae is found in seawater, it is halophilic (“salt-loving”), so it can be cultivated in nutrient media with a higher salt concentration. In addition, it is alkali-tolerant and can therefore grow in nutrient media with an alkaline pH value .

V. cholerae operates a chemoorganotrophic and heterotrophic metabolism , it uses organic compounds as an energy source and also to build up cellular substances. Its metabolism is similar to that of the Enterobacteriaceae representatives , it can utilize several substrates in one fermentation . Various carbohydrates (e.g. glucose , sucrose and mannose ) are broken down into acids and other products by fermentation. Gas is not formed in the process. It also has the enzymes ornithine decarboxylase (ODC) and lysine decarboxylase (LDC), which enable the splitting off of carbon dioxide (CO 2 ) from the amino acids ornithine and lysine . Therefore, a “colorful series” , which is used to differentiate the Enterobacteriaceae, can also be used for the determination of V. cholerae .

genetics

Schematic representation of a bacterial cell with a bacterial chromosome (1) and plasmids (2); Vibrio cholerae contains two bacterial chromosomes, the smaller of which was originally a plasmid.

The genome of the strain Vibrio cholerae O1 N16961 (also known as Biovar El Tor ) was completely sequenced in 2000 . The bacterial strain used for the study was isolated from a stool sample from a patient in a 1971 cholera epidemic in Bangladesh. The genome has a size of 4033 kilobase pairs (kb), which is roughly comparable to the genome size of Escherichia coli . 3887 proteins are annotated . The results of the sequencing of the El-Tor strain and other strains of the species show a GC content (the proportion of the nucleobases guanine and cytosine ) in the bacterial DNA of 47-48 mol percent. This is comparable to the GC content in the DNA of E. coli and other Enterobacteriaceae, which, like Vibrio, belong to the class of Gammaproteobacteria .

The genome of V. cholerae is distributed over two circular chromosomes, which is unusual for bacteria because most bacteria only have a single covalently closed, circular bacterial chromosome. Chromosome 1 of V. cholerae is 2961 kb, while chromosome 2 is smaller with 1072 kb. Most of the genes for important cell functions, such as replication , transcription , translation of DNA and synthesis of the bacterial cell wall , but also the virulence factors, are located on the large chromosome. The smaller chromosome contains genes that are not typical of the Gammaproteobacteria genome . Rather, these are genes that are typically located on a plasmid . For example, there is an integron here (called an integron island ). This gene segment is used to capture genes from a chromosome or a plasmid, which is important for pathogenicity. Since such integrons are otherwise only contained on a plasmid, their discovery on bacterial chromosome 2 led to the assumption that its origin is a “ megaplasmid ” that was taken up by the original form of a Vibrio species.

proof

Yellow colored colonies of Vibrio cholerae on TCBS agar

The test methods used in food microbiology for Vibrio cholerae and other Vibrio species are certified by ISO 21872 and in the USA by the Bacteriological Analytical Manual (bacteriological-analytical manual, abbreviation BAM) of the Food and Drug Administration (FDA) - the US Food and Drug Safety Authority - mandatory.

For samples such as water or food, a first carried enrichment of V. cholerae in alkaline peptone water . In addition to peptone (a mixture of peptides and amino acids ), this nutrient broth has a high concentration of sodium chloride and an alkaline pH value of 8.5; these two parameters inhibit the growth of numerous other bacteria. If the cultivation takes place in this medium at 42 ° C, the enrichment is even more selective, since the growth of other mesophilic bacteria is inhibited by the high temperature. For the isolation of V. cholerae a small volume of nutrient broth on is after enrichment TCBS agar plated, who as Vibrio - selective medium is used.

For clinical specimens such as B. stool samples or vomit, these can be spread with a swab on the selective medium. However, selection with an alkaline medium such as alkaline peptone water is also recommended here. In addition to the selective medium TCBS agar, a nutrient medium that is not very selective should also be inoculated. The direct detection of mobile vibrions in clinical specimens is also carried out with the aid of dark field microscopy or phase contrast microscopy , although no reliable identification of the species is possible in this way.

TCBS agar stands for Thiosulfate Citrate Bile Sucrose Agar and refers to the most important components of the nutrient medium: High concentrations of sodium thiosulfate and sodium citrate largely inhibit the growth of gram-negative Enterobacteriaceae, while ox bile ( bile in English ) the growth of the gram-positive accompanying flora, especially the enterococci prevented. Sucrose (in English sucrose ) is the only carbohydrate in TCBS agar and restricts the growth of bacteria, which can not utilize sucrose. Together with the pH indicators it contains, the breakdown of carbohydrates by Vibrio species can be made visible through acid formation. If not inoculated, the medium has an alkaline pH value (pH 8.6), which also inhibits the growth of other bacteria.

Colonies grown on TCBS agar must be examined further in order to differentiate between the different Vibrio species. Biochemical tests for identification include, as already described, the catalase and oxidase test , as well as typical tests from a "colorful series" , which examine, among other things, the usability of various carbohydrates and other substrates. A rapid determination system based on this in miniature format ( Analytical Profile Index ) for the determination of bacteria from the Enterobacteriaceae and Vibrionaceae families is commercially available.

The assignment to the serotypes can be made by means of an agglutination test. A polyvalent O-specific antiserum is used, which leads to agglutination with sample material which contains O-antigens from V. cholerae . This procedure can be carried out as a so-called latex test (latex agglutination test), in which the antigen-antibody reaction is made visible with the help of latex particles. Detection of V. cholerae may also directly using the ELISA -Verfahrens (quantitative detection of antigens ) are performed. Both the toxin-producing strains of serogroups O1 and O139 and non-toxin-producing strains are recorded. A rectal swab from a patient can be used as a sample, and samples from the environment - for example water samples - can also be examined.

In addition to the detection of the bacterium, there is also evidence of the cholera toxin formed by the pathogen . This is done with the help of the latex agglutination test already mentioned. The antibodies required are obtained from the blood serum of rabbits that have been immunized with the purified cholera toxin. Such test systems are referred to as RPLA ( reversed passive latex agglutination ; translated as "reverse passive latex agglutination") and are commercially available as a simultaneous detection method for cholera toxin and the heat-labile enterotoxin LT from enterotoxic Escherichia coli (ETEC). The test is carried out with microtiter plates , diluted liquid cultures are used as samples, to which the latex suspension is added. The detection limit is 1–2 ng / ml, the sensitivity and the specificity of the RPLA test are comparable to the ELISA method. The detection of certain parts of the bacterial genome using the PCR method ( polymerase chain reaction ) is similarly specific . The aim of the investigation is that part of the genome in which toxin formation is coded . The detection is carried out with the help of the multiplex PCR method, which also enables the simultaneous differentiation from other enterotoxins that cause gastroenteritis .

Occurrence

The Congo is used as a source of drinking water or for washing, which creates the conditions for the transmission of Vibrio cholerae .

Vibrio cholerae is an aquatic bacterium, so it occurs in water, both in fresh water and in sea ​​water , whereby the brackish and coastal waters are of particular importance. The water is also transmitted to humans. Especially untreated or insufficiently treated drinking water is the reason for the transmission, as is food that has come into contact with contaminated water. In Germany, too, V. cholerae was found occasionally in stagnant waters - v. a. at water temperatures> 20 ° C - proven. The vibrios found were not cholera toxin producers.

Vegetable foods can also be contaminated with V. cholerae if faeces containing the pathogen are applied to fields as fertilizer or if the food is wetted with contaminated water to keep it fresh. More often, however, the bacterium is found in animal foods that come from the sea. In patients suffering from cholera, the pathogen can be detected in the stool , vomit and duodenal juice . After the disease has been overcome, V. cholerae can be detected in the stool for a few weeks, but permanent excretors are rare.

Systematics

External system

Vibrio cholerae is a typical representative of the genus Vibrio , which is also known colloquially as Vibrionen. In addition to the cholera pathogen, the species V. parahaemolyticus , V. vulnificus and V. alginolyticus are also of medical importance. In addition to the vibrio there are other species that the family of Vibrionaceae belong, while this is the only family in the order Vibrionales represents.

Internal system

The species includes numerous phyla . If one differentiates Vibrio cholerae according to the O-antigens , over 150 serotypes can be recorded.The serogroups V. cholerae O1 and O139 are considered to be the causative agents of cholera, the serogroup O1 being responsible for most cholera outbreaks. This group includes the classic biotype and the El Tor biotype. Both biotypes each have two own serotypes, which are referred to as Inaba and Ogawa.

For the cholera epidemics in the 19th and 20th centuries, the classic biotype is regarded as the pathogen that causes a significantly more serious course of the disease. Since the 1970s, infections by the biotype (also known as biovar) El Tor have been observed, which are usually less severe. Only in Bangladesh have outbreaks due to the classic biotype been documented since 1982, which is why the southern coastal region of Bangladesh is regarded as a habitat of the classic biotype. The term so-called NAG-Vibrionen (non-agglutinating vibrions), i.e. strains that do not agglutinate with an O-specific antiserum, originates from that time . In fact, the non-agglutination only relates to the antigen O1, which was the only known O-antigen at the time. These non-O1 strains of V. cholerae also cause gastroenteritis , but at the time it was assumed that only the serogroup O1 was involved in epidemics.

That changed in 1992 when several cholera outbreaks were recorded in southern and eastern India and southern Bangladesh, which spread to south-eastern Asia. Strains of serogroup O139 ( V. cholerae O139 synonym Bengal), a previously unknown serovar, were identified as the pathogen . These strains show no agglutination with polyclonal or monoclonal antibodies which are directed against the V. cholerae O1 antigen. Studies from the epidemic area also led to the assumption that immunity acquired against V. cholerae O1 El Tor does not protect against infection with V. cholerae O139. In contrast to the serogroup O1, these Bengal strains have a capsule , and parts of the gene complex responsible for the O1 antigen show a modified structure or are completely absent. In addition to the serogroups V. cholerae O1 and O139, there are also so-called non-O1 and non-O139 strains, which also cause diseases, but which are not described as an outbreak of cholera.

etymology

Otto Friedrich Müller , a Danish naturalist, developed the first attempts to classify and describe microorganisms in 1786 and coined the term "trembling animals" or vibrions for mobile microorganisms ( vibrare from Latin means "rapidly moving back and forth", "vibrating"). The species name refers to the disease cholera caused by the bacterium , comes from ancient Greek and means "river of yellow bile". The biotype El Tor is named after the place El Tor on the Sinai Peninsula , where the German bacteriologist Felix Gotschlich isolated the biotype in a quarantine station in 1905. The patients in the quarantine station were pilgrims who had returned from Mecca. World icon

Medical importance

Pathogenicity

The pathogenicity of Vibrio cholerae is based on the one hand on the release of an exotoxin that acts as an enterotoxin on the intestine. This so-called cholera toxin (CTX) is responsible for the symptoms of the infectious disease cholera . The formation of cholera toxin is coded in the genome of the bacterium, but this only applies to pathogenic strains such as the serogroups V. cholerae O1 and O139.

In the bacterial chromosome can be found on a so-called pathogenicity genes containing various virulence factors encode, in V. cholerae these pathogenicity is called VPI ( V . Cholerae P athogenicity I denotes SLAND). The VPI contains the tcpA gene for the TCP factor (toxin coregulated pili) , which means that the bacterium forms special pili (type IV pili). These are surface structures made of proteins that resemble the flagella , but unlike these, they do not serve for active movement . These filamentous cell surface structures act as adhesins and thus enable the bacterial cells to adhere to the surface of the microvilli of the intestinal cells. So they are another virulence factor.

It is believed that the genes of the pathogenicity island come from a bacteriophage (also called phage for short). In a study from 1999 it was shown that the tcp genes of the VPI are identical to those of a phage, which is consequently referred to as VPIΦ (VPIphi, Vibrio Pathogenicity Island Phage). VPIΦ is therefore responsible for the horizontal gene transfer between V. cholerae strains. Furthermore, the TcpA subunit of the type IV pilus was recognized as the envelope protein of the bacteriophage VPIΦ.

A bacteriophage injects its DNA (shown in blue) into a bacterial cell , the genetic material is integrated into the bacterial chromosome and is now called a prophage .

The type IV pilus also serves as a receptor for another bacteriophage called CTX bezeichnet (CTXphi, cholera toxin phage). The ctxAB genes, which code for the subunits A and B of the cholera toxin, are found in its genome . CTXΦ is a temperate phage that integrates into the bacterial chromosome. The DNA of the phage is built into the chromosome of the bacterium - this is called prophage - and with each subsequent cell division the genes of the prophage and those of the bacterium are duplicated and passed on. As a result, the bacterium forms the accessory colony factor (ACF) and the cholera toxin is released. Usually this so-called lysogenic cycle remains , so there is no activation of the lytic cycle , the phages do not lyse the cell, and so CTXΦ remains in the cell as a prophage and multiplies with it.

Only if a V. cholerae strain has both virulence factors can it be regarded as a pathogenic strain. In a genetic investigation of 300 non-pathogenic strains, a VPI with a tcpA gene could only be detected in 15 strains, of which CTXΦ could also be detected as a prophage in nine strains. None of the strains examined had only the ctxAB genes, which confirms that the phage CTXΦ only integrates into the bacterial chromosome if the tcpA gene is already present in the VPI. The strains investigated which show VPI and CTXΦ as prophages belong to different serogroups than O1 and O139 and are not involved in cholera epidemics. This is justified by deviations in the DNA sequence of the VPI or the prophages. It is concluded from this that these non-epidemic strains evolved from non-pathogenic V. cholerae that incorporated VPI and CTXΦ into their genome by horizontal gene transfer.

The origin of the ctxAB genes by the phage CTXΦ has already been proven, so a transfer of the genes between V. cholerae strains (ie within a species) could be shown with the help of the CTXΦ. This horizontal gene transfer - caused by a phage - is then referred to as transduction . Transmission between species is also possible, for example from V. cholerae to Vibrio mimicus . Furthermore, the VPI of V. mimicus has an identical DNA sequence with that of V. cholerae O1 N16961 in individual genes , which is explained as horizontal gene transfer by the phage VPIΦ.

The origin of the genes in the pathogenicity island has not yet been conclusively clarified. Normally, certain stimuli can be used to reactivate a prophage in such a way that its genes are "read". In the case of temperate phages, this leads to the lysogenic cycle becoming a lytic cycle in which the phages are subsequently released. Treatment of the prophage of V. cholerae with mitomycin C or with UV radiation , an takes place induction of the lytic cycle. In a study based on this from 2003, the production of extracellular CTXΦ could be detected in both V. cholerae O139 strains and in the biovar El Tor. However, this did not succeed for the phage VPIΦ. This does not conclusively prove whether the horizontal gene transfer of the VPI is actually based on a phage or on another mechanism.

Regardless of the serogroup present, V. cholerae is assigned to risk group 2 by the Biological Agents Ordinance in conjunction with the TRBA ( Technical Rules for Biological Agents) 466 .

Sources of infection

The infection of Vibrio cholerae is always orally , in most cases, the recording is done through contaminated drinking water , partly on foods. Direct fecal-oral transmission from person to person is rare. The pathogen can also be detected on plankton components and freshwater algae . They then contaminate fish and seafood that, raw or undercooked, can infect the person who eats these foods. The bacterium can survive for several weeks if there is enough moisture. However, studies on volunteers have shown that a fairly large amount of V. cholerae must be ingested in order to cause cholera infection. The gastric acid reduces the number of pathogens, so that the inoculum must contain about 10 8 to 10 9 bacteria. If V. cholerae is ingested with food, the infectious dose is much lower, in this case the intake of 10 4 to 10 6 bacteria is sufficient . In the case of hypoacidity - if, for example, there is less stomach acid due to the effect of medication - the infectious dose is only 10 3 to 10 4 pathogens. If they get into the small intestine , they can reproduce there again because of the prevailing alkaline environment.

Infectious diseases

Vibrio cholerae attaches to the epithelial cells of the small intestine, multiplies there and releases an exotoxin that acts as an enterotoxin . The result is gastroenteritis with very severe diarrhea ("rice water stool") which, if left untreated, can lead to desiccosis (dehydration of the body) with loss of electrolytes and thus death. Infections caused by the El Tor biotype are usually easier. In addition, there are mild courses without typical symptoms , in which the pathogen can still be detected.

Some of the Non-O1 and Non-O139 strains occasionally lead to gastroenteritis which is vaguely similar to the symptoms of cholera. Studies of such strains have shown that the permeability factor they produce differs immunologically from the cholera toxin. Others of the Non-O1 and Non-O139 strains produce a heat-stable exotoxin that has led to septic infection in patients with certain pre-existing conditions. The main difference is that none of these strains release cholera toxin.

In Germany, direct or indirect evidence of Vibrio cholerae must be reported by name in accordance with Section 7 of the Infection Protection Act . According to § 6 IfSG, suspicion of illness as well as cholera and death must be reported by name; according to § 42 , the person concerned is prohibited from working in certain food establishments.

In Austria, according to Section 1 (1) of the Epidemic Act 1950, cholera is notifiable with regard to suspected illnesses and deaths.

In Switzerland, the positive laboratory analysis findings to be Vibrio cholerae notifiable and that after the Epidemics Act (EpG) in connection with the epidemic Regulation and Annex 3 of the Regulation of EDI on the reporting of observations of communicable diseases of man .

Therapy and prevention

Therapy with antibiotics such as ciprofloxacin can shorten the course of the disease, but is not sufficient on its own. At the same time, replacement of fluids and electrolytes must be provided, as their losses in the course of cholera, if left untreated, lead to a mortality rate of 60%. There is also a vaccine available against cholera in which an inactivated vaccine (inactivated cells of V. cholerae is used), the orally as oral vaccine is administered.

Food microbiological importance

For drinking water as an important food in force in Germany drinking water regulations . According to this, the drinking water must not contain pathogens "in concentrations that cause harm to human health" (Section 5 (1) TrinkwV 2001). The water treatment in the production of drinking water protection in Europe from attack by Vibrio cholerae . In countries or regions with low hygiene standards - caused in particular by inadequate sewage disposal - the occurrence of the cholera pathogen in drinking water or water used for human consumption is a danger. The Robert Koch Institute recommends only boiled water or mineral water prepacked for use dishwashing drinking, brushing teeth or when traveling to these regions. A vaccination for travelers is not generally recommended by the Robert Koch Institute (with reference to the World Health Organization WHO).

Plate of raw seafood

V. cholerae can also be transmitted via fish and seafood . The German Society for Hygiene and Microbiology (DGHM e.V.) therefore recommends that sea fish from warmer regions be subjected to a food microbiological test for vibrions. If pathogenic species are detected, further studies of the toxin production capacity are necessary. The pathogen is also found on plankton components and freshwater algae and can therefore contaminate fish and seafood. Therefore one should avoid raw or insufficiently cooked foods of this kind for prophylaxis. This applies in particular to areas in which the pathogen is likely to occur. You should also avoid raw salads as they could be washed with contaminated water. The rule “Boil it, cook it, peel it, or forget it!” (“Cook it, cook it, peel it or forget it!”) Is a guideline for cholera prophylaxis when traveling to endangered areas.

A study from 1995 showed that many foods, when intentionally contaminated with the pathogen, allow the bacteria to grow or at least to persist, in some cases up to three months. The foods examined included yogurt , milk , jam , salad , pasta and sausages . The vibrions remained viable, especially when the pH of the food in question was neutral or only slightly acidic .

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literature

  • Michael T. Madigan, John M. Martinko, Jack Parker: Brock Microbiology. German translation edited by Werner Goebel, 1st edition. Spektrum Akademischer Verlag GmbH, Heidelberg / Berlin 2000, ISBN 978-3-8274-0566-1 .
  • Herbert Hof, Rüdiger Dörries: Dual Series: Medical Microbiology . 3. Edition. Thieme Verlag, Stuttgart 2005, ISBN 978-3-13-125313-2 .
  • Helmut Hahn, Stefan HE Kaufmann, Thomas F. Schulz, Sebastian Suerbaum (eds.): Medical microbiology and infectiology . 6th edition. Springer Verlag, Heidelberg 2009, ISBN 978-3-540-46359-7 .
  • I. Kaye Wachsmuth, Paul A. Blake, Orjan Olsvik (eds.): Vibrio cholerae and cholera: Molecular to global perspectives . 1st edition. ASM Press, Washington 1994, ISBN 1-55581-067-5 .

Individual evidence

  1. ^ M. Bentivoglio, P. Pacini: Filippo Pacini: a determined observer. In: Brain research bulletin. Volume 38, Number 2, 1995, pp. 161-165, doi: 10.1016 / 0361-9230 (95) 00083-Q . PMID 7583342 .
  2. ↑ Real Academia de la Historia (ed.): Joaquín Balcells y Pasqual , 2018, archive link (Spanish)
  3. ↑ Col·legi Oficial de Metges de Barcelona (ed.): Joaquim Balcells i Pascual , 2015, archive link ( Catalan )
  4. ↑ Norman Howard-Jones: Robert Koch and the cholera vibrio: a centenary. In: British medical journal (Clinical research ed.). Volume 288, Number 6414, February 1984, pp. 379-381, PMID 6419937 . PMC 1444283 (free full text).
  5. ↑ a b Taxonomy Browser Vibrio cholerae. In: National Center for Biotechnology Information (NCBI) website . Retrieved March 8, 2013 .
  6. ↑ a b c Vibrio cholerae O1 bv El Tor, N16961. In: Website Genomes Online Database (GOLD) . Retrieved March 8, 2013 .
  7. ↑ a b Vibrio cholerae El Tor N16961 Genome Page. (No longer available online.) In: J. Craig Venter Institute (JCVI) website . Archived from the original on February 23, 2014 ; Retrieved March 8, 2013 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.  @1@ 2Template: Webachiv / IABot / cmr.jcvi.org
  8. ↑ a b c John F. Heidelberg, Jonathan A. Eisen u. a .: DNA sequence of both chromosomes of the cholera pathogen Vibrio cholerae. In: Nature. Volume 406, number 6795, August 2000, pp. 477-483, doi: 10.1038 / 35020000 . PMID 10952301 .
  9. ↑ a b c d e f David KR Karaolis, Sita Somara, David R. Maneval, Judith A. Johnson, James B. Kaper: A bacteriophage encoding a pathogenicity island, a type-IV pilus and a phage receptor in cholera bacteria. In: Nature. Volume 399, number 6734, May 1999, pp. 375-379, doi: 10.1038 / 20715 . PMID 10360577 .
  10. ^ A b Hans G. Schlegel, Christiane Zaborosch: General microbiology . 7th edition. Thieme Verlag, Stuttgart / New York 1992, ISBN 3-13-444607-3 , p. 24, 65, 117 .
  11. ↑ a b JA Johnson, CA Salles et al. a .: Vibrio cholerae O139 synonym bengal is closely related to Vibrio cholerae El Tor but has important differences. In: Infection and Immunity. Vol. 62, Number 5, May 1994, pp. 2108-2110, PMID 8168977 . PMC 186475 (free full text).
  12. ↑ Michael T. Madigan, John M. Martinko, Jack Parker: Brock Microbiology. German translation edited by Werner Goebel, 1st edition. Spektrum Akademischer Verlag GmbH, Heidelberg / Berlin 2000, ISBN 978-3-8274-0566-1 , pp. 536-538.
  13. ↑ a b c d e Charles A. Kaysner, Angelo DePaola, Jr .: Bacteriological Analytical Manual, chapter 9: Vibrio. In: Website of the Food and Drug Administration (FDA) . Retrieved March 18, 2013 .
  14. ↑ a b c d e f g h i j k l Profiles of rare and imported infectious diseases. (PDF) (No longer available online.) In: Website of the Robert Koch Institute (RKI). September 15, 2011, pp. 61-62 , archived from the original on December 30, 2013 ; Retrieved November 6, 2013 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.  @1@ 2Template: Webachiv / IABot / www.rki.de
  15. ↑ a b c d e Herbert Hof, Rüdiger Dörries: Dual series: Medical microbiology . 3. Edition. Thieme Verlag, Stuttgart 2005, ISBN 978-3-13-125313-2 , p. 400-404 .
  16. ↑ Vibrio cholerae. In: National Center for Biotechnology Information (NCBI) Genome website . Retrieved November 1, 2013 .
  17. ↑ Michael T. Madigan, John M. Martinko, Jack Parker: Brock Microbiology. German translation edited by Werner Goebel, 1st edition. Spectrum Akademischer Verlag GmbH, Heidelberg / Berlin 2000, ISBN 978-3-8274-0566-1 , p. 534.
  18. ^ Rapid testing solutions for the detection of Vibrio cholerae. (No longer available online.) In: Merck Millipore website . Archived from the original on November 3, 2013 ; Retrieved November 2, 2013 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.  @1@ 2Template: Webachiv / IABot / www.merckmillipore.de
  19. ↑ Data sheet ENDO agar for microbiology (PDF) from Merck , accessed on November 2, 2013.
  20. ↑ F. Burkhardt: The bacteriological diagnosis of the Vibrio EI Tor infection. In: Zentralblatt für Bakteriologie, Parasitenkunde, Infectious Diseases and Hygiene. 1. Dept. of medical-hygienic bacteriology, virus research and parasitology. Originals. Volume 212, Number 1, December 1969, pp. 177-189, PMID 4195371 .
  21. ↑ Data sheet TCBS-Agar (Vibrio Selective Agar) (PDF) from Merck , accessed on November 2, 2013.
  22. ^ ID 32 biochemical identification (rapid ID 32 E); Vibrionaceae, Enterobacteriaceae. (No longer available online.) In: website of bioMérieux Deutschland GmbH . Archived from the original on January 5, 2014 ; Retrieved March 4, 2013 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.  @1@ 2Template: Webachiv / IABot / www.biomerieux.de
  23. ↑ U. Tuteja, S. Kumar et al. a .: Simultaneous direct detection of toxigenic and non-toxigenic Vibrio cholerae from rectal swabs and environmental samples by sandwich ELISA. In: Journal of medical microbiology. Volume 56, Number 10, October 2007, pp. 1340-1345, doi: 10.1099 / jmm.0.47166-0 . PMID 17893171 .
  24. ↑ DL Evers, J. He, JT Mason, TJ O'Leary: The liposome PCR assay is more sensitive than the Vibrio cholerae enterotoxin and Escherichia coli heat-labile enterotoxin reversed passive latex agglutination test at detecting cholera toxin in feces and water. In: Journal of clinical microbiology. Volume 48, Number 12, December 2010, pp. 4620-4622, doi: 10.1128 / JCM.02019-10 . PMID 20962142 . PMC 3008463 (free full text).
  25. ↑ Oxoid ™ VET-RPLA Toxin Detection Kit. In: thermofisher.com. Thermo Fisher Scientific , accessed January 9, 2017 .
  26. ↑ RJ Almeida, FW Hickman-Brenner a. a .: Comparison of a latex agglutination assay and an enzyme-linked immunosorbent assay for detecting cholera toxin. In: Journal of clinical microbiology. Volume 28, Number 1, January 1990, pp. 128-130, PMID 2298870 . PMC 269552 (free full text).
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  28. ↑ a b c d Michael T. Madigan, John M. Martinko, Jack Parker: Brock Mikrobiologie. German translation edited by Werner Goebel, 1st edition. Spectrum Akademischer Verlag GmbH, Heidelberg / Berlin 2000, ISBN 978-3-8274-0566-1 , pp. 1098-1099.
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  34. ↑ Michael T. Madigan, John M. Martinko, Jack Parker: Brock Microbiology. German translation edited by Werner Goebel, 1st edition. Spektrum Akademischer Verlag GmbH, Heidelberg / Berlin 2000, ISBN 978-3-8274-0566-1 , p. 95.
  35. ↑ SM McLeod, HH Kimsey, BM Davis, MK Waldor: CTXphi and Vibrio cholerae: exploring a newly recognized type of phage-host cell relationship. In: Molecular microbiology. Volume 57, Number 2, July 2005, pp. 347-356, doi: 10.1111 / j.1365-2958.2005.04676.x . PMID 15978069 .
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  43. ^ Text of the Drinking Water Ordinance (TrinkwV 2001) from juris. Retrieved March 24, 2013.
  44. ↑ Specialist group for food microbiology and hygiene, working group for microbiological guidelines and warning values ​​of the DGHM e. V .: Microbiological guideline and warning values ​​for the assessment of food (as of May 2012), Chapter 21 Guideline and warning values ​​for sea fish. (No longer available online.) In: Website of the German Society for Hygiene and Microbiology (DGHM) . Archived from the original on February 11, 2013 ; Retrieved March 24, 2013 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.  @1@ 2Template: Webachiv / IABot / www.dghm.org
  45. ↑ Frequently asked questions and information for travelers. (PDF; 25 kB) In: Health topics: Cholera on the World Health Organization (WHO) website . Retrieved March 24, 2013 .

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