Vibrio cholerae
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''Vibrio cholerae'' is a species of
Gram-negative Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
,
facultative anaerobe A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococc ...
and comma-shaped
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
. The bacteria naturally live in
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estu ...
or
saltwater Saline water (more commonly known as salt water) is water that contains a high concentration of dissolved salts (mainly sodium chloride). On the United States Geological Survey (USGS) salinity scale, saline water is saltier than brackish water, ...
where they attach themselves easily to the chitin-containing shells of crabs, shrimps, and other shellfish. Some strains of ''V. cholerae'' are pathogenic to humans and cause a deadly disease
cholera Cholera is an infection of the small intestine by some strains of the bacterium ''Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea that lasts a few days. Vomiting and ...
, which can be derived from the consumption of undercooked or raw marine life species. ''V. cholerae'' was first described by Félix-Archimède Pouchet in 1849 as some kind of protozoa.
Filippo Pacini Filippo Pacini (25 May 1812 – 9 July 1883) was an Italian anatomist, posthumously famous for isolating the cholera bacterium ''Vibrio cholerae'' in 1854, well before Robert Koch's more widely accepted discoveries 30 years later. Pacini was born ...
correctly identified it as a bacterium and from him, the scientific name is adopted. The bacterium as the cause of cholera was discovered by
Robert Koch Heinrich Hermann Robert Koch ( , ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera (though the bacteri ...
in 1884.
Sambhu Nath De Sambhunath De ; (1 February 1915 – 15 April 1985) was an Indian medical scientist and researcher, who discovered the cholera toxin, the ''animal model of cholera'', and successfully demonstrated the method of transmission of cholera pathogen ...
isolated the cholera toxin and demonstrated the toxin as the cause of cholera in 1959. The bacterium has a
flagellum 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 ...
at one pole and several pili throughout its cell surface. It undergoes respiratory and fermentative metabolism. Two serogroups called O1 and O139 are responsible for cholera outbreaks. Infection is mainly through drinking contaminated water, therefore is linked to sanitation and hygiene. When ingested, it invades the intestinal mucosa can cause diarrhea and vomiting in a host within several hours to 2–3 days of ingestion. Oral rehydration solution and antibiotics such as
fluoroquinolones A quinolone antibiotic is a member of a large group of broad-spectrum bacteriocidals that share a bicyclic core structure related to the substance 4-quinolone. They are used in human and veterinary medicine to treat bacterial infections, as we ...
and
tetracycline Tetracycline, sold under various brand names, is an oral antibiotic in the tetracyclines family of medications, used to treat a number of infections, including acne, cholera, brucellosis, plague, malaria, and syphilis. Common side effects in ...
s are the common treatment methods. ''V. cholerae'' has two circular DNA. One DNA produces the
cholera toxin Cholera toxin (also known as choleragen and sometimes abbreviated to CTX, Ctx or CT) is AB5 multimeric protein complex secreted by the bacterium ''Vibrio cholerae''. CTX is responsible for the massive, watery diarrhea characteristic of cholera i ...
(CT), a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
that causes profuse, watery
diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...
(known as "rice-water stool"). But the DNA does not directly code for the toxin as the genes for cholera toxin are carried by CTXphi (CTXφ), a
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacteri ...
(virus). The virus only produces the toxin when inserted into the bacterial DNA.
Quorum sensing In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...
in ''V. cholerae'' is well studied and it activates host immune signaling and prolongs host survival, by limiting the bacterial intake of nutrients, such as
tryptophan Tryptophan (symbol Trp or W) is an α-amino acid that is used in the biosynthesis of proteins. Tryptophan contains an α-amino group, an α- carboxylic acid group, and a side chain indole, making it a polar molecule with a non-polar aromatic ...
, which further is converted to serotonin. As such, quorum sensing allows a
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
interaction between host and pathogenic bacteria.


Discovery


Initial observations

During the third global pandemic of cholera (1852–1859), there were several scientific research to understand the etiology of the disease. The miasma theory, which posited that infections spread through contaminated air, was no longer a satisfactory explanation. An English physician John Snow was the first to give convincing evidence in London in 1854 that cholera was spread from drinking water – a contagion, not miasma. Yet he could not identify the pathogens, which made most people still believe in the miasma origin. ''V. cholerae'' was first observed and recognized under microscope by a French zoologist Félix-Archimède Pouchet. In 1849, Pouchet examined the stool samples of four people having cholera. His presentation before the French Academy of Sciences on 23 April was recorded as: " ouchetcould verify that there existed in these holera patientsdejecta an immense quantity of microscopic
infusoria Infusoria are minute freshwater life forms including ciliates, euglenoids, protozoa, unicellular algae and small invertebrates. Some authors (e.g., Bütschli) used the term as a synonym for Ciliophora. In modern formal classifications, the term ...
." But he made a mistake in believing that the organisms were infusoria, a name then used for microscopic protists, thereby attributing them as ''Vibrio rigula,'' a species of protozoan described by Danish naturalist Otto Friedrich Müller in 1786.


Identification of the bacterium

An Italian physician,
Filippo Pacini Filippo Pacini (25 May 1812 – 9 July 1883) was an Italian anatomist, posthumously famous for isolating the cholera bacterium ''Vibrio cholerae'' in 1854, well before Robert Koch's more widely accepted discoveries 30 years later. Pacini was born ...
, while investigating cholera outbreak in Florence in the late 1854, identified the causative pathogen as a new type of bacterium. He performed autopsies of corpses and made meticulous microscopic examinations of the tissues and body fluids. From feces and
intestinal mucosa The gastrointestinal wall of the gastrointestinal tract is made up of four layers of specialised tissue. From the inner cavity of the gut (the lumen) outwards, these are: # Mucosa # Submucosa # Muscular layer # Serosa or adventitia The muco ...
, he identified many comma-shaped bacilli. Reporting his discovery before the Società Medico-Fisica Fiorentina (Medico-Physician Society of Florence) on 10 December, and published in the 12 December issue of the ''Gazzetta Medica Italiana'' (''Medical Gazette of Italy''), Pacini stated:Pacini thus introduced the name ''vibrioni'' (Latin ''vībro'' means "to move rapidly to and fro, to shake, to agitate"). A
Catalan Catalan may refer to: Catalonia From, or related to Catalonia: * Catalan language, a Romance language * Catalans, an ethnic group formed by the people from, or with origins in, Northern or southern Catalonia Places * 13178 Catalan, asteroid #1 ...
physician Joaquim Balcells i Pascual also reported such bacterium around the same time. The discovery of the new bacterium was not regarded as medically important as the bacterium was not directly attributed to cholera. Pacini also stated that there was no reason to say that the bacterium caused the disease since he failed to make pure culture and perform experiment, which was necessary 'to attribute the quality of contagion to cholera'. The miasma theory was still not ruled out.


Rediscovery

The medical importance and relationship of the bacterium and cholera was discovered by a German physician
Robert Koch Heinrich Hermann Robert Koch ( , ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera (though the bacteri ...
. In August 1883, Koch, with a team of German physicians, went to Alexandria, Egypt, to investigate cholera epidemic there. Koch found that the intestinal mucosa of people who died of cholera always had the bacterium, yet could not confirm if it was the causative agent. He moved to Calcutta (now Kolkata) India, where the epidemic was more severe. It was from here that he isolated the bacterium in pure culture on 7 January 1884. He subsequently confirmed that the bacterium was a new species, and described as "a little bent, like a comma." He reported his discovery to the German Secretary of State for the Interior on 2 February, and published in the ''Deutsche Medizinische Wochenschrift'' (''German Medical Weekly''). Although Koch was convinced that the bacterium was the cholera pathogen, he could not entirely establish a critical evidence the bacterium produced the symptoms in healthy subjects (an important element in what was later known
Koch's postulates Koch's postulates ( )"Koch"
''
Sambhu Nath De Sambhunath De ; (1 February 1915 – 15 April 1985) was an Indian medical scientist and researcher, who discovered the cholera toxin, the ''animal model of cholera'', and successfully demonstrated the method of transmission of cholera pathogen ...
in Calcutta isolated the cholera toxin and showed that it caused cholera in healthy subjects that the bacterium-cholera relationship was fully proven.


Taxonomy

Pacini had used the name "''vibrio cholera''", without proper
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms * Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition ...
rendering, for the name of the bacterium. Following Koch's description, a scientific name ''Bacillus comma'' was popularised. But an Italian bacteriologist Vittore Trevisan published in 1884 that Koch's bacterium was the same as that of Pacini's and introduced the name ''Bacillus cholerae.'' A German physician
Richard Friedrich Johannes Pfeiffer Richard Friedrich Johannes Pfeiffer FRS (27 March 1858 – 15 September 1945) was a German physician and bacteriologist. Pfeiffer was born to Otto Pfeiffer, a German pastor of the local Evangelical parish, and Natalia née Jüttner, in Treus ...
renamed it as ''Vibrio cholerae'' in 1896. The named was adopted by the Committee of the Society of American Bacteriologists on Characterization and Classification of Bacterial Types in 1920. In 1964, Rudolph Hugh of the George Washington University School of Medicine proposed to use the genus ''Vibrio'' with the type species ''V. cholerae'' (Pacini 1854) as a permanent name of the bacterium, regardless of the same name for protozoa. It was accepted by the Judicial Commission of the International Committee on Bacteriological Nomenclature in 1965, and the International Association of Microbiological Societies in 1966.


Characteristics

''V. cholerae'' is a highly motile, comma shaped, gram-negative rod. The active movement of ''V. cholerae'' inspired the genus name because "vibrio" in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
means "to quiver". Except for v.cholerae and v.mimicus, all other vibrio species are
halophilic The halophiles, named after the Greek word for "salt-loving", are extremophiles that thrive in high salt concentrations. While most halophiles are classified into the domain Archaea, there are also bacterial halophiles and some eukaryotic species, ...
. Initial isolates are slightly curved, whereas they can appear as straight rods upon laboratory culturing. The bacterium has a
flagellum 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 ...
at one cell pole as well as pili. It tolerates alkaline media that kill most intestinal commensals, but they are sensitive to acid. It is a
facultative anaerobe A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococc ...
, and can undergo respiratory and fermentative metabolism. It measures 0.3 μm in diameter and 1.3 μm in length with average swimming velocity of around 75.4 μm/sec.


Pathogenicity

''V. cholerae''
pathogenicity In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ge ...
genes code for proteins directly or indirectly involved in the virulence of the bacteria. To adapt the host intestinal environment and to avoid being attacked by bile acids and antimicrobial peptides, ''V. cholera'' uses its outer membrane vesicles (OMVs). Upon entry, the bacteria sheds its OMVs, containing all the membrane modifications that make it vulnerable for the host attack. During infection, ''V. cholerae'' secretes
cholera toxin Cholera toxin (also known as choleragen and sometimes abbreviated to CTX, Ctx or CT) is AB5 multimeric protein complex secreted by the bacterium ''Vibrio cholerae''. CTX is responsible for the massive, watery diarrhea characteristic of cholera i ...
(CT), a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
that causes profuse, watery
diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...
(known as "rice-water stool"). This cholera toxin contains 5 B subunits that plays a role in attaching to the intestinal epithelial cells and 1 A subunit that plays a role in toxin activity''.'' Colonization of the small intestine also requires the toxin coregulated
pilus A pilus (Latin for 'hair'; plural: ''pili'') is a hair-like appendage found on the surface of many bacteria and archaea. The terms ''pilus'' and '' fimbria'' (Latin for 'fringe'; plural: ''fimbriae'') can be used interchangeably, although some r ...
(TCP), a thin, flexible, filamentous appendage on the surface of bacterial cells. Expression of both CT and TCP is mediated by two component systems (TCS), which typically consist of a membrane-bound
histidine kinase Histidine kinases (HK) are multifunctional, and in non-animal kingdoms, typically transmembrane, proteins of the transferase class of enzymes that play a role in signal transduction across the cellular membrane. The vast majority of HKs are hom ...
and an intracellular response element. TCS enable bacteria to respond to changing environments. In ''V. cholerae'' several TCS have been identified to be important in colonization, biofilm production and virulence.
Small RNA Small RNA (sRNA) are polymeric RNA molecules that are less than 200 nucleotides in length, and are usually non-coding. RNA silencing is often a function of these molecules, with the most common and well-studied example being RNA interference (RN ...
s (sRNA) have been identified as targets of ''V. cholerae'' TCS. Here, sRNA molecules bind to mRNA to block translation or induce degradation of inhibitors of expression of virulence or colonization genes. In ''V. cholerae'' the TCS EnvZ/OmpR alters gene expression via the sRNA ''coaR'' in response to changes in
osmolarity Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). The osmolarity of a solution is usually expressed as Osm/L ...
and pH. An important target of ''coaR'' is ''tcpI'', which negatively regulates expression of the major subunit of the TCP encoding gene (''tcpA''). When ''tcpI'' is bound by ''coaR'' it is no longer able to repress expression ''tcpA'', leading to an increased colonization ability. Expression of ''coaR'' is upregulated by EnvZ/OmpR at a pH of 6,5, which is the normal pH of the intestinal lumen, but is low at higher pH values. ''V. cholerae'' in the intestinal lumen utilizes the TCP to attach to the intestinal mucosa, not invading the mucosa. After doing so it secretes cholerae toxin causing its symptoms. This then increases cyclic AMP or cAMP by binding (cholerae toxin) to adenylyl cyclase activating the GS pathway which leads to efflux of water and sodium into the intestinal lumen causing watery stools or rice watery stools. ''V. cholerae'' can cause syndromes ranging from asymptomatic to cholera gravis. In
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
areas, 75% of cases are asymptomatic, 20% are mild to moderate, and 2-5% are severe forms such as cholera gravis. Symptoms include abrupt onset of watery
diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...
(a grey and cloudy liquid), occasional
vomiting Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
, and abdominal cramps.
Dehydration In physiology, dehydration is a lack of total body water, with an accompanying disruption of metabolic processes. It occurs when free water loss exceeds free water intake, usually due to exercise, disease, or high environmental temperature. Mil ...
ensues, with symptoms and signs such as thirst, dry mucous membranes, decreased skin turgor, sunken eyes,
hypotension Hypotension is low blood pressure. Blood pressure is the force of blood pushing against the walls of the arteries as the heart pumps out blood. Blood pressure is indicated by two numbers, the systolic blood pressure (the top number) and the dia ...
, weak or absent
radial pulse In human anatomy, the radial artery is the main artery of the lateral aspect of the forearm. Structure The radial artery arises from the bifurcation of the brachial artery in the antecubital fossa. It runs distally on the anterior part of the ...
,
tachycardia Tachycardia, also called tachyarrhythmia, is a heart rate that exceeds the normal resting rate. In general, a resting heart rate over 100 beats per minute is accepted as tachycardia in adults. Heart rates above the resting rate may be normal ( ...
,
tachypnea Tachypnea, also spelt tachypnoea, is a respiratory rate greater than normal, resulting in abnormally rapid and shallow breathing. In adult humans at rest, any respiratory rate of 1220 per minute is considered clinically normal, with tachypnea b ...
, hoarse voice,
oliguria Oliguria or hypouresis is the low output of urine specifically more than 80 ml/day but less than 400ml/day.Boon et al, Davidson's Principles & Practice of Medicine (20th Ed), p475 The decreased output of urine may be a sign of dehydration, kidney ...
, cramps,
kidney failure Kidney failure, also known as end-stage kidney disease, is a medical condition in which the kidneys can no longer adequately filter waste products from the blood, functioning at less than 15% of normal levels. Kidney failure is classified as eit ...
,
seizure An epileptic seizure, informally known as a seizure, is a period of symptoms due to abnormally excessive or synchronous neuronal activity in the brain. Outward effects vary from uncontrolled shaking movements involving much of the body with l ...
s,
somnolence Somnolence (alternatively sleepiness or drowsiness) is a state of strong desire for sleep, or sleeping for unusually long periods (compare hypersomnia). It has distinct meanings and causes. It can refer to the usual state preceding falling asleep ...
, coma, and death. Death due to dehydration can occur in a few hours to days in untreated children. The disease is also particularly dangerous for pregnant women and their fetuses during late pregnancy, as it may cause premature labor and fetal death. A study done by the Centers for Disease Control (CDC) in Haiti found that in pregnant women who contracted the disease, 16% of 900 women had fetal death. Risk factors for these deaths include: third trimester, younger maternal age, severe dehydration, and vomiting Dehydration poses the biggest health risk to pregnant women in countries that there are high rates of cholera. In cases of cholera gravis involving severe dehydration, up to 60% of patients can die; however, less than 1% of cases treated with rehydration therapy are fatal. The disease typically lasts 4–6 days. Worldwide,
diarrhoeal disease Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery defecation, bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration of ...
, caused by cholera and many other pathogens, is the second-leading cause of death for children under the age of 5 and at least 120,000 deaths are estimated to be caused by cholera each year. In 2002, the WHO deemed that the case fatality ratio for cholera was about 3.95%.


Cholera illness and symptoms

''V. cholerae'' infects the intestine and causes diarrhea, the hallmark symptom of cholera. Infection can be spread by eating contaminated food or drinking contaminated water. It also can spread through skin contact with contaminated human feces. Not all infection indicate symptoms, only about 1 in 10 people develop diarrhea. The major symptoms include: watery diarrhea, vomiting, rapid heart rate, loss of skin elasticity, low blood pressure, thirst, and muscle cramps. This illness can get serious as it can progress to kidney failure and possible coma. If diagnosed, it can be treated using medications.


Disease occurrence

''V. cholerae'' has an endemic or epidemic occurrence. In countries where the disease has been for the past three years and the cases confirmed are local (within the confines of the country) transmission is considered to be "endemic." Alternatively, an outbreak is declared when the occurrence of disease exceeds the normal occurrence for any given time or location."World Health Organization, Disease Outbreaks." World Health Organization, World Health Organization, 8 Mar. 2016, www.searo.who.int/topics/disease_outbreaks/en/. Epidemics can last several days or over a span of years. Additionally, countries that have an occurrence of an epidemic can also be endemic. The longest standing ''V. chloerae'' epidemic was recorded in Yemen. Yemen had two outbreaks, the first occurred between September 2016 and April 2017, and the second began later in April 2017 and recently was considered to be resolved in 2019."Mystery of Yemen Cholera Epidemic Solved." ScienceDaily, ScienceDaily, 2 Jan. 2019, www.sciencedaily.com/releases/2019/01/190102140745.htm. The epidemic in Yemen took over 2,500 lives and impacted over 1 million people of Yemen. More outbreaks have occurred in Africa, the Americas, and Haiti.


Preventive measures

When visiting areas with epidemic cholera, the following precautions should be observed: drink and use bottled water; frequently wash hands with soap and safe water; use chemical toilets or bury feces if no restroom is available; do not defecate in any body of water and cook food thoroughly. Supplying proper, safe water is important. A precaution to take is to properly sanitize. Hand hygiene is an essential in areas where soap and water is not available. When there is no sanitation available for hand washing, scrub hands with ash or sand and rinse with clean water. A single dose
vaccine A vaccine is a biological Dosage form, preparation that provides active acquired immunity to a particular infectious disease, infectious or cancer, malignant disease. The safety and effectiveness of vaccines has been widely studied and verifie ...
is available for those traveling to an area where cholera is common. There is a ''V. cholerae'' vaccine available to prevent disease spread. The vaccine is known as the, "oral cholera vaccine" (OCV). There are three types of OCV available for prevention: Dukoral®, Shanchol™, and Euvichol-Plus®. All three OCVs require two doses to be fully effective. Countries who are endemic or have an epidemic status are eligible to receive the vaccine based on several criteria: Risk of cholera, Severity of cholera, WASH conditions and capacity to improve, Healthcare conditions and capacity to improve, Capacity to implement OCV campaigns, Capacity to conduct M&E activities, Commitment at national and local level"Oral Cholera Vaccines." World Health Organization, World Health Organization, 17 May 2018, www.who.int/cholera/vaccines/en/. Since May the start of the OCV program to May 2018 over 25 million vaccines have been distributed to countries who meet the above criteria.


Treatment

The basic, overall treatment for Cholera is re-hydration, to replace the fluids that have been lost. Those with mild dehydration can be treated orally with an oral rehydration solution (ORS). When patients are severely dehydrated and unable to take in the proper amount of ORS, IV fluid treatment is generally pursued. Antibiotics are used in some cases, typically
fluoroquinolones A quinolone antibiotic is a member of a large group of broad-spectrum bacteriocidals that share a bicyclic core structure related to the substance 4-quinolone. They are used in human and veterinary medicine to treat bacterial infections, as we ...
and
tetracycline Tetracycline, sold under various brand names, is an oral antibiotic in the tetracyclines family of medications, used to treat a number of infections, including acne, cholera, brucellosis, plague, malaria, and syphilis. Common side effects in ...
s.


Genome

''V. cholerae'' has two circular
chromosomes A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
, together totalling 4 million
base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
of DNA sequence and 3,885 predicted
genes In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
. The genes for cholera toxin are carried by CTXphi (CTXφ), a
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacteri ...
inserted into the ''V. cholerae'' genome. CTXφ can transmit cholera toxin genes from one ''V. cholerae'' strain to another, one form of
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). H ...
. The genes for toxin coregulated pilus are coded by the Vibrio
pathogenicity island Pathogenicity islands (PAIs), as termed in 1990, are a distinct class of genomic islands acquired by microorganisms through horizontal gene transfer. Pathogenicity islands are found in both animal and plant pathogens. Additionally, PAIs are found i ...
(VPI). The entire genome of the virulent strain'' V. cholerae'' El Tor N16961 has been sequenced, and contains two circular chromosomes. Chromosome 1 has 2,961,149
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
s with 2,770
open reading frame In molecular biology, open reading frames (ORFs) are defined as spans of DNA sequence between the start and stop codons. Usually, this is considered within a studied region of a prokaryotic DNA sequence, where only one of the six possible readin ...
s (ORF's) and chromosome 2 has 1,072,315 base pairs, 1,115 ORF's. The larger first chromosome contains the crucial genes for toxicity, regulation of toxicity, and important cellular functions, such as
transcription Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including: Genetics * Transcription (biology), the copying of DNA into RNA, the fir ...
and
translation Translation is the communication of the meaning of a source-language text by means of an equivalent target-language text. The English language draws a terminological distinction (which does not exist in every language) between ''transla ...
. The second chromosome is determined to be different from a
plasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria; how ...
or
megaplasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria; howe ...
due to the inclusion of housekeeping and other essential genes in the genome, including essential genes for metabolism, heat-shock proteins, and 16S rRNA genes, which are ribosomal subunit genes used to track evolutionary relationships between bacteria. Also relevant in determining if the replicon is a chromosome is whether it represents a significant percentage of the genome, and chromosome 2 is 40% by size of the entire genome. And, unlike plasmids, chromosomes are not self-transmissible. However, the second chromosome may have once been a megaplasmid because it contains some genes usually found on plasmids. ''V. cholerae'' contains a genomic island of
pathogenicity In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ge ...
and is lysogenized with
phage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacter ...
DNA. That means that the genes of a virus were integrated into the bacterial genome and made the bacteria pathogenic. The molecular pathway involved in expression of virulence is discussed in the pathology and current research sections below.


Bacteriophage CTXφ

CTXφ (also called CTXphi) is a
filamentous phage Filamentous bacteriophage is a family of viruses (''Inoviridae'') that infect bacteria. The phages are named for their filamentous shape, a worm-like chain (long, thin and flexible, reminiscent of a length of cooked spaghetti), about 6 nm ...
that contains the genes for
cholera toxin Cholera toxin (also known as choleragen and sometimes abbreviated to CTX, Ctx or CT) is AB5 multimeric protein complex secreted by the bacterium ''Vibrio cholerae''. CTX is responsible for the massive, watery diarrhea characteristic of cholera i ...
. Infectious CTXφ particles are produced when ''V. cholerae'' infects humans. Phage particles are secreted from bacterial cells without
lysis Lysis ( ) is the breaking down of the membrane of a cell, often by viral, enzymic, or osmotic (that is, "lytic" ) mechanisms that compromise its integrity. A fluid containing the contents of lysed cells is called a ''lysate''. In molecular bio ...
. When CTXφ infects ''V. cholerae'' cells, it integrates into specific sites on either chromosome. These sites often contain
tandem array Tandemly arrayed genes (TAGs) are a gene cluster created by tandem duplications, a process in which one gene is duplicated and the copy is found adjacent to the original. They serve to encode large numbers of genes at a time. TAGs represent a la ...
s of integrated CTXφ prophage. In addition to the ''ctxA'' and ''ctxB'' genes encoding cholera toxin, CTXφ contains eight genes involved in phage reproduction, packaging, secretion, integration, and regulation. The CTXφ genome is 6.9 kb long.


Ecology and epidemiology

The main reservoirs of ''V. cholerae'' are aquatic sources such as
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of wate ...
s,
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
s, and
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
, often in association with copepods or other
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
,
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater envir ...
, and aquatic plants. Cholera infections are most commonly acquired from drinking water in which ''V. cholerae'' is found naturally or into which it has been introduced from the feces of an infected person. Cholera is most likely to be found and spread in places with inadequate water treatment, poor sanitation, and inadequate hygiene. Other common vehicles include raw or undercooked fish and shellfish. Transmission from person to person is very unlikely, and casual contact with an infected person is not a risk for becoming ill.''V. cholerae'' thrives in an
aquatic environment An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The ...
, particularly in surface water. The primary connection between humans and pathogenic strains is through water, particularly in economically reduced areas that do not have good water purification systems. Nonpathogenic strains are also present in water ecologies. The wide variety of pathogenic and nonpathogenic strains that co-exist in aquatic environments are thought to allow for so many genetic varieties.
Gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HG ...
is fairly common amongst bacteria, and recombination of different ''V. cholerae'' genes can lead to new virulent strains. A symbiotic relationship between ''V. cholerae'' and ''Ruminococcus obeum'' has been determined. ''R. obeum'' autoinducer represses the expression of several ''V. cholerae''
virulence factors Virulence factors (preferably known as pathogenicity factors or effectors in plant science) are cellular structures, molecules and regulatory systems that enable microbial pathogens (bacteria, viruses, fungi, and protozoa) to achieve the followin ...
. This inhibitory mechanism is likely to be present in other gut microbiota species which opens the way to mine the gut microbiota of members in specific communities which may utilize autoinducers or other mechanisms in order to restrict colonization by ''V. cholerae'' or other enteropathogens. Outbreaks of Cholera cause an estimated 120,000 deaths annually worldwide. There has been roughly seven pandemics since 1817, the first. These pandemics first arose in the Indian subcontinent and spread.


Diversity and evolution

Two serogroups of ''V. cholerae'', O1 and O139, cause outbreaks of cholera. O1 causes the majority of outbreaks, while O139 – first identified in
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
in 1992 – is confined to Southeast Asia. Many other serogroups of ''V. cholerae'', with or without the cholera toxin gene (including the nontoxigenic strains of the O1 and O139 serogroups), can cause a cholera-like illness. Only toxigenic strains of serogroups O1 and O139 have caused widespread epidemics. ''V. cholerae'' O1 has two biotypes, classical and
El Tor El Tor is a particular strain of the bacterium ''Vibrio cholerae'', the causative agent of cholera. Also known as ''V. cholerae'' biotype eltor, it has been the dominant strain in the seventh global cholera pandemic. It is distinguished from t ...
, and each biotype has two distinct serotypes, Inaba and Ogawa. The symptoms of infection are indistinguishable, although more people infected with the El Tor biotype remain asymptomatic or have only a mild illness. In recent years, infections with the classical biotype of ''V. cholerae'' O1 have become rare and are limited to parts of Bangladesh and
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. Recently, new variant strains have been detected in several parts of Asia and Africa. Observations suggest these strains cause more severe cholera with higher case fatality rates.


Natural genetic transformation

''V. cholerae'' can be induced to become competent for natural genetic transformation when grown on chitin, a biopolymer that is abundant in aquatic habitats (e.g. from crustacean exoskeletons). Natural genetic transformation is a sexual process involving DNA transfer from one bacterial cell to another through the intervening medium, and the integration of the donor sequence into the recipient genome by
homologous recombination Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may ...
. Transformation competence in ''V. cholerae'' is stimulated by increasing cell density accompanied by nutrient limitation, a decline in growth rate, or stress. The ''V. cholerae'' uptake machinery involves a competence-induced
pilus A pilus (Latin for 'hair'; plural: ''pili'') is a hair-like appendage found on the surface of many bacteria and archaea. The terms ''pilus'' and '' fimbria'' (Latin for 'fringe'; plural: ''fimbriae'') can be used interchangeably, although some r ...
, and a conserved DNA binding protein that acts as a ratchet to reel DNA into the cytoplasm. There are two models of genetic transformation, sex hypothesis and competent bacteria.


Gallery

File:Vibrio diagram.png, Diagram of the bacterium, ''V. cholerae'' File:Vibrio cholerae on TCBS agar.jpg, Yellow colored (sucrose-fermenting) colonies of ''Vibrio cholerae'' on TCBS agar. File:Vibrio cholerae.jpg, Transmission electron microscope image of ''Vibrio cholerae'' that has been negatively stained.


See also

*
Drinking water Drinking water is water that is used in drink or food preparation; potable water is water that is safe to be used as drinking water. The amount of drinking water required to maintain good health varies, and depends on physical activity level, a ...
* Haiti cholera outbreak *
Cholera vaccine Cholera vaccines are vaccines that are effective at preventing cholera. For the first six months after vaccination they provide about 85percent protection, which decreases to 50percent or 62percent during the first year. After two years the level ...


References


External links


Copepods and cholera in untreated water

Vibrio cholerae El Tor N16961 Genome Page
* mepage
Type strain of ''Vibrio cholerae'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Authority control Vibrionales Gram-negative bacteria Cholera Bacteria described in 1854