This article was originally published in the Encyclopedia of Food and Health published by Elsevier, and the

attached copy is provided by Elsevier for the author's benefit and for the benefit of the authors institution, for
non-commercial research and educational use including without limitation use in instruction at your institution,
sending it to specific colleagues who you know, and providing a copy to your institutions administrator.

All other uses, reproduction and distribution, including without limitation commercial reprints, selling or
licensing copies or access, or posting on open internet sites, your personal or institutions website or repository,
are prohibited. For exceptions, permission may be sought for such use through Elsevier's permissions site at:

http://www.elsevier.com/locate/permissionusematerial

Suffredini E., and Caburlotto G. (2016) Vibrio: Types, Properties, and Determination. In: Caballero, B., Finglas,
P., and Toldr, F. (eds.) The Encyclopedia of Food and Health vol. 5, pp. 413-417. Oxford: Academic Press.
2016 Elsevier Ltd. All rights reserved.

Author's personal copy

Vibrio: Types, Properties, and Determination

E Suffredini, Istituto Superiore di Sanita`, Rome, Italy
G Caburlotto, Istituto Zooprofilattico Sperimentale delle Venezie, Adria, Italy
2016 Elsevier Ltd. All rights reserved.

Vibrio Types and Properties

Vibrio Genus
The genus Vibrio belongs to the Vibrionaceae family and
includes bacteria autochthonous of the marine environment.
To date, the genus includes more than 90 species and its
taxonomy is continuously updated due to the addition of
new species. Morphologically, vibrios are Gram-negative
straight or curved rods, with a size between 0.5 and 0.8 mm in
width and 23 mm in length; they are motile by means of a
single polar flagellum and display both respiratory metabolism
and fermentative metabolism. Vibrios are nonsporogenous
microorganisms, but survival in the environment is facilitated
by resistance forms as the viable but nonculturable state
(VBNC), a dormant state characterized by a reduction of cells
size, and a shift to alternative metabolic pathways. The genus
Vibrio includes both halotolerant and halophilic bacteria: halotolerant vibrios are capable of growth in the absence of salt
(NaCl), whereas halophilic vibrios are dependent on the presence of NaCl for multiplication. Due to their flexible physiology, vibrios are one of the most common microorganisms in
marine and estuarine waters throughout the world, and they
can persist in a variety of aquatic ecosystems, including freshwater habitats (V. cholerae and V. mimicus) and seawater, both
as free-living cells and in association with aquatic animals and
plants.
Several Vibrio species are known to be associated with outbreaks and sporadic cases due to the consumption of contaminated water, food manipulated in inadequate hygiene
conditions, seafood (raw fish and shellfish), or the exposure
of skin lesion (cuts, open wounds, or abrasions) to aquatic
environments and marine animals (Table 1). Pathogenicity
toward humans is commonly associated with the presence of
specific genetic features (e.g. toxin-coding genes and virulence
factors) whose variable distribution among the strains often
results in the coexistence of pathogenic and nonpathogenic
populations in the same species. The most commonly known
pathogen is V. cholerae, the etiologic agent of cholera, one of
the few foodborne illnesses with pandemic potential. Other
relevant species that may cause disease, mainly by seafood
ingestion, are V. parahaemolyticus and V. vulnificus, which are
normal inhabitants of marine environments and are isolated
from seawater and seafood with frequencies varying, according
to the studies, from less than 10% up to 36% for finfish
samples and 100% for shellfish. Variations in Vibrio prevalence
and levels are indeed expected, in relation to the food type
(with higher presence in filter-feeding animals as shellfish), the
geographic origin of the seafood product, and the sampling
period. Vibrio levels in the aquatic environment, in fact, are
known to correlate with water temperatures, and detection is
therefore more likely in warm months than in winter periods.
Unfortunately, because vibrios are part of the normal marine

Encyclopedia of Food and Health

microflora and, with the exception of V. cholerae, are not associated with fecal pollution or anthropic impact on seafood
production or harvesting areas, control measures can only
rely on adequate manipulation of the products (rapid refrigeration, uninterrupted cold chain, and proper cooking) and on
the monitoring of the environmental conditions that may
favor the increase of Vibrio levels.
Further to the three major human pathogens (V. cholerae,
V. parahaemolyticus, and V. vulnificus) and to the other species
(V. alginolyticus, V. fluvialis, V. furnissi, V. metschnikovii, etc.)
that are occasionally responsible for disease in susceptible
individuals, several Vibrio species (e.g., V. anguillarum,
V. harveyi, and V. salmonicida) express pathogenicity toward
marine animals (fish, shellfish, or mammals) with considerable economic loss in the aquaculture sector due to large
mortality episodes or to prolonged decrease of production.
Significantly, some Vibrio species display potential for both
severe diseases in humans and in aquatic animals, as is the
case of V. vulnificus, which causes mortality in eels and deadly
wound infections in aquaculture professionals (fishermen,
processors, etc.), or of V. parahaemolyticus, causative agent of
gastroenteritis in seafood consumers and responsible for the
early mortality syndrome in shrimps.

Vibrio cholerae
V. cholerae is the etiologic agent of cholera, an infectious disease responsible, in areas endemic for this microorganism, for
large epidemics and for a high number of deaths, especially in
the presence of inadequate hygiene standards and improper
treatment of wastewaters. The first description of a Vibrio-like
microorganism from patients affected by cholera dates back to
1854 with Filippo Pacini and first isolation of V. cholerae was
achieved in 1884 by Robert Koch. The modern history of
cholera is marked by seven pandemics, during which the disease spread at a global level with reiterated and geographically
shifting epidemic peaks.
V. cholerae includes more than 200 hundred serogroups
according to somatic antigen O, a thermostable polysaccharide
of the cellular wall. Serotypes O1 and O139 are associated with
cholera epidemics, whereas the other serotypes, generally
defined as V. cholerae non-O1/non-O139, are occasionally
responsible for sporadic cases of gastroenteritis, mild forms
of disease similar to cholera, or extraintestinal infections. Serogroup O1 can be further divided in two serotypes (Ogawa and
Inaba) and two distinct biotypes: the biotype Classic, deemed
responsible for the first six pandemics (18171923), and the
biotype El Tor, so called by the name of the quarantine center
where it was first isolated, responsible for the seventh ongoing
pandemic (1961). Biotype differentiation was based initially
on the characteristic hemolytic effect of El Tor variant on sheep
erythrocytes, but this feature became progressively less discriminating during the seventh pandemics as the nonhemolytic

http://dx.doi.org/10.1016/B978-0-12-384947-2.00724-8

The Encyclopedia of Food and Health, (2016), vol. 5, pp. 413-417

413

Author's personal copy

414

Vibrio: Types, Properties, and Determination

Table 1
diseases

Most relevant Vibrio species associated with human

Species

Source

Disease

V. alginolyticus

Seafood,
seawater
Water or food

Wound infection, ear

infections, gastroenteritis
Gastroenteritis

Water or food

Gastroenteritis, wound
infection, ear infection
Gastroenteritis
Gastroenteritis
Gastroenteritis, ear infection
Primary septicemia,
gastroenteritis
Gastroenteritis, ear infection,
wound infection
Gastroenteritis
Primary septicemia, wound
infection, bacteremia,
gastroenteritis

V. cholerae
O1-O139
V. cholerae non-O1/
non-O139
V. fluvialis
V. furnissii
V. hollisae
V. metschnikovii
V. mimicus
V. parahaemolyticus
V. vulnificus

Seafood
Seafood
Shellfish
Shellfish,
seawater
Shellfish,
seawater
Shellfish
Shellfish,
seawater

strains overcame the hemolytic ones. The distinction, however,

maintained an epidemiological convenience, as the different
biotypes seem to correlate with the clinical severity of the
disease, being the El Tor type less aggressive (only 2% of
patients develop severe disease, compared to 11% of the classical infection). Until 1992, the distinction between V. cholerae
O1 and non-O1 (also described as NAG, nonagglutinable
V. cholerae) was considered pivotal to differentiate between
strains with potential for epidemic cholera and nonpathogenic
V. cholerae isolates. In early 1993, however, reports appeared in
India and Bangladesh of a severe cholera-like disease associated with a new serogroup, O139, sometimes referred to as
V. cholerae Bengal, for the origin of the strain. Since its first
appearance, this serogroup, characterized by hybrid features of
O1 and non-O1 strains, has spread to other countries (China,
Thailand, Pakistan, Nepal, and Malaysia) and, although
remaining geographically confined to Southeast Asia, has compelled a global revision of serological testing.
As other Vibrio species, V. cholerae is part of the autochthonous bacterial flora of estuarine areas, and its growth may
occur in variable temperature conditions (from 10 to 43  C)
and in NaCl concentrations ranging from 0.1% to 4.0%. Halotolerance of V. cholerae is a key feature of its transmission,
especially in developing countries, where ingestion of contaminated water is probably the most important vehicle. Free in the
drinkable water, V. cholerae can, in fact, survive for several days
(up to 13) and can contribute to persistence of the infection
also via food prepared with contaminated water, as in the case
of some outbreaks generated by rice dishes or fruit juices, or
with the use of contaminated water for the irrigation of vegetables. It is also demonstrated that V. cholerae can persist in the
environment in the absence of human disease, possibly
through the colonization of the surface of zooplankton (e.g.,
copepods) and aquatic plants. Another relevant feature for
V. cholerae transmission is related to its pH tolerance.
V. cholerae may grow in a pH range between 5.0 and 9.6 with
pH 7.6 being the optimum pH and displays sensitivity to acidic
environment (which, in the form of gastric acids, provides the

major human defense barrier to Vibrio infection). Foods

involved in V. cholerae transmission (crustaceans, shellfish,
cooked meat and rice dishes, nonalcoholic and noncarbonated
drinks, and raw vegetables and fruits) usually display neutral or
nearly neutral pH, and thus, this aspect can be taken into
account to orientate the investigations in outbreak settings.
The main virulence factor of V. cholerae is represented by the
cholera toxin (CT), a typical AB subunit toxin encoded by the ctx
gene (ctxA and ctxB), which is present on a horizontally transmissible filamentous bacteriophage (CTXF). CT is responsible
for the osmotic disequilibrium in intestinal lumen at the origin
of the massive water flows (rice water feces) characteristic of
cholera disease. CT, however, is not the only pathogenicity
determinant of V. cholerae, as multiple systems are involved in
regulation of virulence. Among these, the ToxR regulon controls
the expression of several (at least 17) genes involved in the
pathogenic mechanism of V. cholerae, and therefore, toxR gene
is often used as a molecular marker in V. cholerae analysis.
Further to CT, other factors are expressed in virulent strains:
the zonula occludens toxin (zot), which affects the structure of
tight junction and increases the permeability of the intestinal
mucosa; the accessory CT (ace), which acts on fluid
accumulation; and the toxin-coregulated pilus (TCP), long filaments on the surface of the microorganism with an important
role in adherence and colonization of intestinal mucosa. Also,
non-O1/non-O139 V. cholerae strains may occasionally encode
for some of these genes, as well as for other specific virulence
factors as a heat-stable toxin (codified by the st gene), thermostable direct hemolysins, or shigella-like toxins.

Vibrio parahaemolyticus
V. parahaemolyticus was first described as the etiologic agent of
the shirasu food poisoning in Japan. Late in 1950, a severe
foodborne outbreak (272 cases and 20 fatalities) was reported
in Osaka, with patients suffering from acute gastroenteritis,
severe abdominal pain, vomiting, and diarrhea. Epidemiological investigation led to identify the source of the outbreak to a
small, semidried fish (shirasu) and to the isolation by Dr.
Tsunesaburo Fujino of a new microorganism, Pasteurella parahaemolytica, later renamed Vibrio parahaemolyticus.
Since then, V. parahaemolyticus outbreaks have been repeatedly reported in the world, and at present, V. parahaemolyticus is
considered the first cause of the nonviral infections related to
shellfish consumption in the United States, Japan, and Southeast Asia. This microorganism, in fact, displays the features
common to Vibrio genus, as the natural occurrence in the
marine environment and the correlation with water temperature and salinity, and is therefore commonly present in seafood products, with densities usually lower than 103 cfu g 1,
although such levels may be exceeded in products harvested
from warmer seawaters. In addition, V. parahaemolyticus is also
characterized by a remarkable replication time (as short as
89 min in optimal conditions and in the range of
1218 min in seafood) that allows multiplication of the microorganism and rapid reaching of the estimated infectious dose
(105107 cfu for pathogenic strains) in raw, undercooked, or
cross contaminated seafood products that are incorrectly
stored.
As for V. cholerae, not all V. parahaemolyticus strains are
pathogenic. A strong correlation between human disease and

The Encyclopedia of Food and Health, (2016), vol. 5, pp. 413-417

Author's personal copy

Vibrio: Types, Properties, and Determination

production of two toxins (the thermostable direct hemolysin

(TDH) and the TDH-related hemolysin (TRH)) has been demonstrated. TDH production, which induces beta-hemolytic
activity in human erythrocytes on Wagatsuma agar (a process
designated as Kanagawa phenomenon), is dependent upon the
presence and expression of one or more nonidentical copies of
the tdh gene (five variants have been reported), while TRH is
encoded by the trh gene, in which at least two major subgroups
(trh1 and trh2) can be defined. Besides these hemolysins, other
putative virulence factors are currently under investigation, as
it has been shown that about 10% of clinical strains contain
neither tdh nor trh. In particular, the role of the type III secretion systems (T3SSs) detected in V. parahaemolyticus (T3SS2-a
and T3SS2-b) and of its genomic islands (Vibrio pathogenicity
islands (VPaIs)) has been scrutinized since the emergence in
early 1996 of a O3:K6 strain, whose diffusion, first in Asia
(India, Bangladesh, Indonesia, Thailand, Laos, Taiwan, Korea,
and Japan) and then in Europe, Mozambique, the United
States, Mexico, and South American countries, has given rise
to the first V. parahaemolyticus pandemic. Serotyping of
V. parahaemolyticus is based on both somatic O (13 antigens)
and capsular polysaccharide K antigens (65), but up to the
description of the pandemic strain, there was no correlation
between serotype and virulence and, until now, factors providing advantage to O3:K6 over other strains remain unclear.

Vibrio vulnificus
V. vulnificus is the most clinically serious Vibrio infection in
developed countries, accounting in the United States for
almost the totality (95%) of the deaths associated with seafood
consumption. V. vulnificus is responsible, in fact, for severe
infections in susceptible individuals, with two distinct disease
syndromes, primary septicemia (with a case-fatality rate of
approximately 50%) and necrotizing wound infections.
Based upon the variation of their biochemical profiles,
three V. vulnificus biotypes can be distinguished. Strains
belonging to biotype 1 are most commonly associated with
human disease and can be isolated from seafood products
(finfish and bivalve mollusks), biotype 2 is usually considered
as an eel pathogen, while strains belonging to biotype 3, isolated in Israel in 1996, affect humans through wound infection
and bacteremia. V. vulnificus is a widespread inhabitant of
estuarine environments; its presence has been reported in seawater, shellfish, crabs, etc., and a strong correlation has been
detected between water temperature and its isolation.
Infections are usually sporadic, and in almost all of them, the
patient has an underlying chronic disease (e.g., cirrhosis or liveror blood-related disorders) leading to elevated serum iron levels;
other cases are reported in the presence of pathologies as chronic
renal or gastric diseases, immunosuppression, and diabetes.
Many studies have attempted discrimination of virulent and
avirulent strains by comparison of clinical and environmental
strains, but due to the high degree of genetic diversity, a definitive indicator has yet to be identified. V. vulnificus invasivity is
related to the high variety of virulence factors expressed by this
species: the polysaccharide capsule, which is produced by nearly
all strains and confers an opaque aspect to V. vulnificus colonies,
is essential for infection as it protects the microorganism from
phagocytosis and from the activity of the complement system;
the endotoxic lipopolysaccharides is responsible for the typical

415

symptoms (fever, hemorrhage, hypotension, etc.) of V. vulnificus

septicemia; siderophore production allows iron uptake from
host serum; and a large number of extracellular enzymes,
including proteases, elastase, and a hemolysin/cytotoxin with
strong cytolytic activity, support the tissue disruption activity
during the infection.

Other Vibrio Species

Besides the three species previously described, other vibrios
have been recognized as potential pathogens for humans.
V. alginolyticus is one of the Vibrio species most commonly
detected in seawater and in seafood products; infections (otitis
and wound infections) are mostly related to contact with the
marine environment and few reports of its implication as a
foodborne pathogen are available, even though severe cases
have been described in patients with underlying medical conditions. V. mimicus, formerly known as sucrose-negative
V. cholerae non-O1 for its similarity to the more renowned
species, is causative agent of sporadic cases of gastroenteritis,
mainly in relation to consumption of raw shellfish and crustaceans. Most of the virulence factors expressed by V. mimicus are
toxins described in other species, including CT, TDH, zot, and
st of non-O1/non-O139 V. cholerae strains. Finally, two closely
related species, V. fluvialis and V. furnissi, have also been implicated in gastroenteritis episodes or outbreaks; V. fluvialis, in
particular, has been reported in a large number of pediatric
patients in India, and in surveillance data from the Center for
Disease Control and Prevention, it represents approximately
5% of the vibriosis in the United States.

Vibrio Detection
Cultural Methods
In routine analysis of food products, the detection of pathogenic vibrios mostly relies on conventional cultural methods,
followed by biochemical and, for V. cholerae, serological testing. Such approach, applied in internationally recognized
methods as ISO/TS 21872 and the FDA Bacteriological Analytical Manual, includes a preenrichment step in a nonselective
medium (usually alkaline peptone water with an appropriate
concentration of NaCl) and provides resuscitation of bacteria,
including cells in a VBNC state and cells stressed due to product refrigeration or freezing. Enrichment is then followed by
isolation of colonies on selective agar media, with thiosulfate
citrate bile salt agar (TCBS) being the most commonly used.
Random selection of colonies with characteristic features represents a critical point of the procedure, as species of interest in
food analysis (V. cholerae, V. parahaemolyticus, and V. vulnificus)
may be overgrown both during enrichment and on agar plates
by other vibrios as V. alginolyticus or by other genera, therefore
making isolation of relevant colonies less efficient. In this
context, the formulation of new selective and differential
media (e.g., chromogenic media) to use in association with
TCBS has progressively improved the chance to differentiate
the species targeted by the analysis. The isolation of colonies is
critical also under another prospective: as pathogenic strains of
Vibrio species often represent a small minority of the microbial
flora, the number and the proportion of characteristic colonies

The Encyclopedia of Food and Health, (2016), vol. 5, pp. 413-417

Author's personal copy

416

Vibrio: Types, Properties, and Determination

selected for further testing may significantly affect the probability of detecting the pathogenic strains of a species (e.g.,
isolates belonging to the serogroups O1/O139 of V. cholerae
or tdh-positive/trh-positive V. parahaemolyticus strains). Further
to the isolation step, the total performance of the cultural
methods may be challenged also by the identification of the
isolates; the variability of the biochemical profiles of Vibrio
environmental strains, in fact, may negatively affect the accuracy of the biochemical identification by conventional tests,
dichotomous keys, or miniaturized tests, leading to inconclusive results or to misidentifications.
Conventional cultural methods for Vibrio analysis show,
overall, important characteristics that encourage their routine
adoption, including technical easiness, relatively low costs, and
integration with laboratorys workflows. On the other hand,
however, the cultural analytic process is time-consuming (45
days to obtain the identification of the microorganisms),
labor-intensive, and, lastly, unable to provide a characterization of the virulence factors associated with human illness. To
this aim, a progressive integration of the cultural and molecular analytic methods for Vibrio analysis has been pursued, as
testified by the introduction, also in the aforementioned
methods for official controls, of polymerase chain reaction
(PCR) protocols for sample screening, improvement of species
identification, and characterization of pathogenicity factors.

Molecular Methods
During the past years, nucleic acid detection and amplification
methods have significantly broadened the spectrum of diagnostic tools for the analysis of Vibrio species, and to date, a large
number of both conventional and real-time PCR protocols,
isothermal amplification (LAMP) procedures, and some hybridization methods have been published for the detection, identification, characterization, or enumeration of potentially
pathogenic species. One of the major advantages accorded by
nucleic acid-based techniques is related to the fact that, by
targeting directly genetic sequences responsible for or associated
with virulence of defined Vibrio species, they provide more
reliable information on the risk associated with the presence of
pathogenic vibrios in a product. A second advantage is given by
the substantial reduction of the time required for the analysis,
with detection of potentially pathogenic species completed, for
certain protocols, in 24 h or less. In this sense, the molecular
techniques represent the most suitable analytic approach for
industries and food producers to prevent release on the market
of products containing pathogenic Vibrio species, as well as for
control authorities to provide timely results, therefore ensuring
food safety and preventing outbreaks.
Molecular detection methods for Vibrio have been directed,
depending on the scope of the analysis, to either conserved
region of the genome (species detection or isolates
identification) or genes encoding or associated with virulence
(detection of potentially pathogenic strains). PCR targeting
total V. cholerae, for example, has largely relied on the detection
of conserved and widespread genes as toxR, which is involved
in the regulation cascade of virulence-associated genes; ompW
gene, which encodes an outer membrane protein and is conserved in both clinical and environmental isolates; and hlyA
gene, a nonclassical fragment of the hemolysin gene present in

O1, O139, and non-O1/non-O139 V. cholerae strains. Identification of pathogenic strains has, on the other hand, mainly
targeted genes associated with the CT production, including
ctxA and ctxB, or codifying for other virulence factors as zot, ace,
tcp, or st genes. PCR protocols, however, have also been successfully developed for rapid characterization of strains, for
example, with the identification of V. cholerae O1 epidemic
strains through ctxB allelic discrimination or by using rfb gene
for the molecular differentiation of O1 and O139 isolates.
Compared to V. cholerae, a multiplicity of sequences has
been proposed in the last years for the detection of
V. parahaemolyticus in food samples and seawater or for the
identification of isolates. The 16S ribosomal RNA gene, for
example, was repeatedly evaluated as a possible target for
molecular analysis, but high sequence homology with other
species (e.g., 99.7% with V. alginolyticus) and the presence
of intrastrain polymorphisms have hampered its use. GyrB
gene, encoding the B subunit of DNA gyrase, and fragments
of the hsp (heat shock protein) gene, being characterized
by a lower sequence homology with other Vibrio species,
were also suggested as possible targets for differentiation of
V. parahaemolyticus. Three other targets, however, have been
extensively used and compared, especially for the application
in food analysis: the tlh gene, codifying for a thermolabile
hemolysin (TLH) in V. parahaemolyticus and, at present, widely
used in the United States and Japan; the toxR gene, homologous to the regulator gene of V. cholerae and conserved with
low sequence identity in the whole Vibrionaceae family, whose
application is common in Southeast Asia and Europe; and the
pR72H sequence, a restriction fragment generated by HindIII
enzyme that includes a coding (phosphatidylserine synthase
gene) region and a noncoding region and that appears to be
conserved in V. parahaemolyticus strains. In contrast to methods
for V. parahaemolyticus identification, protocols aiming at the
detection of potentially enteropathogenic strains have
unequivocally targeted the tdh and trh genes, often in multiplex
format with each other or with one of the aforementioned
sequences for species identification.
For V. vulnificus, identification has been mainly based on the
cytotoxinhemolysin gene (vvh), thought to be common to all
biotypes, although potential problems of gene loss associated
with the use of this target have been reported. Alternative genomic loci, as the 16S rRNA gene, have been therefore investigated,
both for identification purposes and to achieve differentiation
between clinical and environmental strains; a 17-base sequence
variation in the 16S rRNA gene, in fact, was shown to have a
statistically significant correlation with clinical isolates. Other
genes and sequences, including the virulence-correlated gene
(vcg), a siderophore-encoding gene (viuB), and the capsular
polysaccharide operon, have demonstrated, however, comparable or higher discriminative power and have been widely
applied for V. vulnificus characterization.
In the analysis of foods in which Vibrio species may be
present, nucleic acid amplification-based methods usually provide results with high specificity and information relevant for
the characterization of pathogenicity potential of microorganisms. Nonetheless, a major drawback of these methods is that,
due to the low concentration of the targets and the high inhibitory effect of food matrices on PCR amplification, sensitivity
levels compatible with food analysis can be achieved only on

The Encyclopedia of Food and Health, (2016), vol. 5, pp. 413-417

Author's personal copy

Vibrio: Types, Properties, and Determination

preenriched samples. As a consequence, even though several

real-time qPCR protocols have been developed for analysis of
vibrios in foods, quantification by molecular methods has in
the end settled for the use of MPN-PCR formats that, although
sensitive, call for a considerable increase of analytic work and
costs. An alternative approach for Vibrio enumeration is
granted by colony hybridization techniques, which combine
growth on nonselective agar media with molecular identification/characterization using specific probes, and a few hybridization protocols have been described, especially for
quantitative analysis of V. parahaemolyticus and V. vulnificus.

See also: Codex Alimentarius.

417

Desmarchelier PM, Wong FYK, and Venkateswaran K (2000) Vibrio. In: Robinson R,
Batt C, and Patel P (eds.) Encyclopedia of food microbiology, pp. 22372258.
San Diego: Academic Press.
Drasar BS and Forrest BD (eds.) (1996) Cholera and the ecology of Vibrio cholerae.
London: Chapman & Hall.
Faruque SM and Nair GB (eds.) (2008) Vibrio cholerae: genomics and molecular
biology. Norfolk: Caister Academic Press.
Oliver JD and Kaper JB (2001) Vibrio species. In: Doyle MP, Beuchat LR, and
Monteville TJ (eds.) Food microbiology: fundamentals and frontiers, 2nd ed.,
pp. 263300. Washington, DC: ASM Press.
Parveen S and Tamplin ML (2013) Vibrio vulnificus, Vibrio parahaemolyticus and
Vibrio cholerae. In: Labbe RG and Garca S (eds.) Guide to foodborne pathogens,
2nd ed., pp. 148176. Oxford: Wiley.
Su Y-C (ed.) (2012) Pathogenic vibrios and food safety. Hauppauge, New York: Nova
Science Publishers.
Thompson FL, Austin B, and Swings JG (eds.) (2006) The biology of vibrios.
Washington, DC: ASM Press.

Further Reading
Relevant Websites
Croci L and Suffredini E (2013) Real-time PCR detection of foodborne pathogenic
Vibrio. In: Rodrguez-Lazaro D (ed.) Real-time PCR in food science: current
technology and applications, pp. 135148. Norfolk: Caister Academic Press.

http://www.fda.gov/Food/FoodScienceResearch/LaboratoryMethods/ucm2006949.htm
FDAs Bacteriological Analytical Manual (BAM).

The Encyclopedia of Food and Health, (2016), vol. 5, pp. 413-417