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safe for human consumption, if they are handled and stored
properly.

Vegetable Products
Vegetables may also be fermented to preserve them or form modified products with characteristic flavors and textures. Fermented
vegetable products include pickles, ripe olives, sauerkraut, doenjang, shoyu, stinky tofu, miso, tempeh, injera, kimchi, koji, natto,
soy sauce, brandy, cider, sake, and vinegar; they are produced
from sources such as beans, grains, cucumbers, lettuce, olives,
cabbage, turnips, fruits, and rice. In general, biogenic amine
content is very low in fresh vegetables; however, the concentration increases during the fermentations process and throughout
storage. Sauerkraut is made from shredded cabbage by fermentation with lactic acid bacteria. In the case of fermented vegetable
products, there does not appear to be a significant correlation
between biogenic amine levels and spoilage unlike the situation

with fermented meat products, and compounds like tyramine,
putrescine, and cadaverine are commonly found in fermented
vegetable products (Potter and Hotchkiss 1998, Kalac et al. 1999,
2000a, 2000b).
The amounts of biogenic amines present in fermented vegetable products also vary and depend on factors such as temperature, salt content, and pH/acidity, as well as starter culture type.
These factors all affect the growth and metabolism of the organism, which influence the formation of the biogenic amines. Low
temperatures generally minimize microbial growth and proliferation, and enzyme activities to reduce biogenic amine formation,
while elevated temperatures tend to do the reverse.
Salt plays an important role in the fermentation of vegetables.
By modulating the Aw levels, salting can influence the texture
of fermented vegetable products. For example, under dry salted
fermentation conditions, product firmness is enhanced with increasing salt content. Although a high salt may suppress the
activity of certain (desirable) bacteria, there are halophilic bacteria that are salt tolerant, and advantage may be taken of this
fact to use halophilic bacteria such as Enterococci to overcome
the growth and proliferation of a moderately salt tolerant Lactobacilli species in fermented vegetable products (Kalac et al.
1999).
Acidity and pH are also important in the fermentation of vegetables. Hitherto, the use of starter cultures helps to exclude
undesirable microorganisms by generating an acidic milieu that
is not conducive for the growth of certain microorganisms. Thus,
the pH level in a product can determine the types of microorganisms that would be dominant in the course of the fermentation
process. For instance, Lactobacillus and Streptococcus species
are acid tolerant, and when they are used in mixed cultures,
species like Leuconostoc mesenteroides would initiate the process and produce an acidic environment, and then be replaced
by L. plantarum till the lactic acid level rises sufficiently to kill
the L. plantarum and be replaced by L. pentoaceticus. L. plantarum is a typical tyramine producer; thus, products formed with
such cultures would be expected to accumulate biogenic amines
(Battcock and Azam-Ali 1998, Suzzi and Gardini 2003).

Fermented vegetable products also accumulate biogenic
amines during storage. For example, the levels of tyramine and

putrescine in sauerkraut increase during storage, and the longer
the storage period, the more of these biogenic amines accumulate (Kalac et al. 2000a, 2000b).

Non-fermented Foods
Biogenic amines occur in non-fermented foods also. Fresh fish
and meats in particular are known to contain biogenic amines
due to their high protein and free amino acid contents and their
high perishability. These food products have also been associated with histamine poisoning, which also attests to the presence
of these compounds. Biogenic amines in non-fermented foods
are also formed by the action of decarboxylating and proteolytic
enzymes exuded by microorganisms naturally present as part of
the native microbial flora of fish and meats, or microorganisms
purposefully added as starter culture, or added through contamination. High levels of biogenic amines in foodstuffs may be
used as indices of undesirable microbial growth and spoilage.

Fish
Biogenic amines in fish are of particular health concern because
they cause histamine poisoning in humans (Sanchez-Guerrero
et al. 1997). Histamine poisoning or scombroid fish poisoning (scombrotoxicosis) is the most widespread form of seafood
poisoning in fish-producing and fish-consuming communities.
Fish in the Scombridae and Scomberesocidae families as well
as non-scombroid fish have both been implicated in biogenic
amine poisoning of humans. These include fish species like bluefish (Pomatomus spp.), bonito (Sarda spp.), mackerel (Scomber
spp.), mahi mahi (Coryphaena spp.), marlin (Makaira spp.),
pilchards (Sardina pilchardus), sardines (Sardinella spp.), saury
(Cololabis saida), sockeye salmon (Oncorhynchus nerka), tuna
(Thunnus spp.), and yellowtail (Seriola lalandii) (Lehane and
Olley 2000, Sarkadi 2009). These fish species tend to have high
levels of free histidine in their tissues, which may undergo decarboxylation to form the toxic histamine. The levels of biogenic
amines in fish tissues vary greatly and are influenced by various

factors including muscle type, the native microflora, harvesting, and postharvest management practices (including handling,
processing, transportation and storage; Veciana-Nogues et al.
1997a, 1997b, Gloria et al. 1999).
Of the biogenic amines found in fish tissues, histamine usually tends to be the most abundant, followed by putrescine and
cadaverine in that order. For example, tuna fish samples stored at
20◦ C produced histamine, cadaverine, and putrescine levels of
3103 µg/g, 44.2 µg/g, and 3.0 µg/g, respectively, after 48 hours
of storage (Veciana-Nogues et al. 1997a, 1997b). Nonetheless,
these biogenic amines may not occur in all fish tissues, as is the
case for tyramine, which is absent in sea bream (Sparus aurata)
but present in anchovies (Engraulis mordax; Koutsoumanis et al.
1999).
The relative amounts of biogenic amines and their distribution
in fish tissues depend to a large extent on the types of microorganisms associated with the raw material and their capacity to


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43 Biogenic Amines in Foods

produce decarboxylases and hydrolytic enzymes like proteases
and lipases. Generally, when the microbial load is high, the biogenic amine levels also tend to be high (Du et al. 2002).
Approximately, one-third of the microorganisms isolated from
spoiled tuna and mackerel are decarboxylase enzyme producers. These include Acinetobacter iwoffi, Aeromonas hydrophila,
Clostridium perfringens, Enterobacter aerogenes, Hafnia alvei,
Morganella morganii, Proteus mirabilis, Proteus vulgaris, Pseudomonas fluorescens, Pseudomonas putrefaciens, and Vibrio alginolyticus, and they are members from both mesophilic and psychrophilic organisms. In this regard, the mesophilic types tend
to play a relatively more significant role in histamine production
and fish flesh deterioration than their psychrophilic counterparts,
and species like Morganella morganii, as well as Enterobacteriaceae, Clostridium, and Lactobacillus are of particular importance in terms of histamine production in fish (Omura et al. 1978,
Taylor 1986, Middlebrooks et al. 1988, Lakshmanan et al. 2002,
Kim et al. 2006).
Environmental factors (e.g., temperature, pH, and salt content)
also play a role in biogenic amines formation and accumulation
in fish tissues, and these factors may be manipulated to slow
down biogenic amine formation.
Fish muscle type also influences the amounts of biogenic
amines that form and/or accumulate in fish tissues. In this connection, red-fleshed fish tend to have higher levels of histamine
in their tissues than their white-fleshed counterparts. This is due
to the relatively higher content of the precursor molecule, histidine, in red-fleshed fish tissues, as well as higher amounts of
carbohydrates and lipids, and a slightly more acidic environment, all conditions that favor the growth and proliferation of
decarboxylase positive microorganisms.
The importance of temperature as a foremost manipulative
factor to limit the formation of biogenic amines in fish tissues is
due to the fact that several of the major histidine-decarboxylasepositive bacteria associated with fish are mesophilic. Thus, a decrease in temperature slows down their growth and metabolism
(Ababouch et al. 1991). For example, icing decreases the initial mesophilic microorganism population while increasing the
lag phase of growth. Nevertheless, decreasing product temperature is not an absolute means of control. This is because certain
psychrophilic bacteria can also produce decarboxylases that can
lead to the formation of biogenic amines, even at proper refrigeration temperatures. Furthermore, the decarboxylase enzymes

released in fish tissues are still functional at low temperatures
although at a reduced rate. Thus, proper postharvest fish management is crucial to control formation and accumulation of
biogenic amines in fish flesh.

Meat
Fresh meats (beef, pork, and poultry) are also non-fermented
foods known to have biogenic amines. Their high proteinaceous
nature makes them prone to proteolysis to form free amino acids
that may subsequently be decarboxylated by microbial enzymes
into the biogenic amines. Spermine and spermidine are believed
to be present in all fresh meats (beef, pork, and poultry) at fairly
constant levels, unlike the other biogenic amines that occur in

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varying amounts. Other predominant biogenic amines in fresh
meats are putrescine and cadaverine, together with relatively
lower levels of histamine, tryptamine, and tyramine (Paulsen
and Bauer 2007, Galgano et al. 2009). For beef, tyramine or a
combination of the three most prevalent biogenic amines, tyramine, putrescine, and cadaverine, have been recommended for
use as index of quality or spoilage (Vinci and Antonelli 2002).
Fresh poultry meat can also have significant amounts of biogenic amines and the accumulation of compounds such as spermidine, putrescine, cadaverine, histamine, and tyramine tend to
increase during storage. In poultry, putrescine and cadaverine
tend to be the predominant biogenic amines and may attain total levels of 50–100 mg/kg when stored over extended periods.
Biogenic amines also accumulate more rapidly in raw poultry
meat because of the higher susceptibility of white muscle fibers
to proteolysis versus the darker muscle fibers of red meats, beef,
or pork (Perez et al. 1998, Balamatsia et al. 2006, Patsias et al.
2006).
Enterobacteriaceae species are believed to be primarily responsible for putrescine, cadaverine, and histamine formation

in poultry meat. Other microorganisms that have been identified
as major biogenic amine producers include Brochothrix thermosphacta, Carnobacterium divergens, Carnobacterium piscicola,
Citrobacter freundii, Enterobacter agglomerans, Escherichia
coli, Escherichia vulnaris, Hafnia alvei, L. curvatus, Morganella
morganii, Proteus alcalifaciens, and Serratia liquifaciens
(Masson et al. 1996, Pircher et al. 2007).
Biogenic amine formation and/or accumulation in nonfermented meats are influenced by essentially the same factors as for fermented meats. For example, the type of meat is
important, as it determines the availability of the substrate or
precursor molecules. Relatively larger quantities of cadaverine
are produced in both red and white meat due to the high levels
of free lysine in both muscle types; however, red meats tend to
have higher tyramine contents from their higher free tyrosine
contents.
In fresh meats, temperature, salt content, and pH all play an
important role in biogenic amine production. However, this attribute varies from species to species. For example, low pHs
enhance the production of histamine, tyramine, and tryptamine
in poultry meats by lactic acid than at a higher pH, while Enterobacteriaceae produced less cadaverine at low pHs. High
salt concentrations also has a greater inhibitory effect on the
formation of biogenic amines, and this is due to inhibition of
microbial growth by the reduced Aw from the high salt content;
higher temperatures promote more biogenic amines formation in
beef, pork, and poultry samples than frozen temperatures. Thus,
a higher salt content or lower storage temperature (or both) is
recommended for curtailing biogenic amine formation and extending the shelf life of meat products (Chander et al. 1989,
Karpas et al. 2002).

SIGNIFICANCE TO THE FOOD INDUSTRY
Biogenic amines are of particular interest to the food industry
due to their toxicity, and their use as indicators of food quality or spoilage (Ruiz-Capillas and Moral 2004). For example,



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histamine poisoning occurs following ingestion of foods that
have large quantities of histamine (Stratton et al. 1991). Scombroid fish if not handled properly can accumulate high quantities
of histamine, and such products can pose a health hazard when
consumed. Other fermented foods, such as cheese, wine, dry
sausage, sauerkraut, miso, and soy sauce have also been linked
with biogenic amines poisoning. Histamine and tyramine toxicities have been extensively investigated, and biogenic amines
such as β-phenylethylamine and tyramine have been linked to
hypertensive crises (Shalaby 1996) and migraines. Other biogenic amines such as the HCAs have also been implicated in the
formation of mutagenic or carcinogenic nitrosamines (Stratton
et al. 1991), and the potential health risks of these compounds
are areas of active research to better understand the extent of the

health hazards posed by these compounds in food products.

Health Effects
Biogenic amines participate in various functions connected with
cell growth and metabolism in humans. For example, spermine
and spermidine are involved in free radicals scavenging, cell
membrane channels modulation, and in DNA synthesis. However, the compounds must be present in minute quantities to
carry out useful functions, because they can be toxic in high
quantities. Other biogenic amines, such as histamine and tyramine, occur naturally in small quantities and participate in allergic and inflammatory responses (Khan et al. 1992, Ha et al.
1998, Schneider et al. 2002). The metabolism of histamine entails absorption by active or passive transport through the lumen of the GIT. During this passage, most of the histamine
is broken down by the enzymes diamine oxidase (DAO) and
histamine-N-methyltransferase (HNMT) into useful intermediates like imidazoleacetic acid and N-methyl histamine. These
molecules together with the residual histamine are transported
in the blood to the liver where they are degraded further. In the
body, histamine binds to receptors to initiate various responses,
including smooth muscle contraction, vasodilation, alteration
of blood pressure, and enhancing vascular permeability. Nevertheless, high levels of histamine are potentially toxic and this
intoxication manifests itself within 2 hours after ingestion and
may last up to 16 hours. The symptoms include allergic-type
reactions and gastrointestinal problems in the form of nausea,
cramps, vomiting, and diarrhea (Izquierdo-Pulido et al. 1999,
Attaran and Probst 2002, Maintz and Novak 2007). Histamine
intoxication can also manifest as neurological and cutaneous
disorders in the form of dizziness, flushing, headache, rash, and
itchy eyes. These symptoms are attributed to the release of nitric
oxide from vascular endothelial cells to bring about vasodilation
(Becker et al. 2001, Predy et al. 2003, Borade et al. 2007). In
rare cases, histamine poisoning could cause severe hypotension,
atrial arrhythmias, and loss of consciousness (McInerney et al.
1996).

Various factors may influence histamine toxicity. These include the use of drugs that inhibit mono- and DAO enzymes,
thereby preventing detoxification of histamine by deamination,
gastrointestinal diseases, alcohol consumption, and bacteria endotoxins. Other biogenic amines like putrescine and cadaverine

can also compete for the same catabolic enzymes as histamine,
thus inhibiting the effectiveness of histamine detoxification and
making the toxic compounds accumulate highly in the plasma.
Although histamine intoxication is the type most commonly
associated with biogenic amines, tyramine also has high toxicity. Ingestion of large quantities of tyramine-containing cheeses,
alcohol, chocolate, and other dairy products can elicit tyramine
intoxication, which manifests as headaches and elevated blood
pressure (Millichap and Yee 2003, Panconesi 2008). Tyramine
is catabolized rapidly in the small intestine by MAOs, and monitoring tyramine ingestion is crucial for individuals on MAOinhibiting drugs as their ingestion could bring on hypertensive
crisis (Schmidt and Ferger 2004, Vlieg-Boerstra et al. 2005).
Tyramine poisoning can also cause nausea, vomiting, and even
death from intracranial hemorrhage. There are also emerging
concerns with nitrosamines because they can form carcinogenic
nitrosamines with cadaverine and putrescine by reacting with nitrites (used to cure meats) or by serving as precursor molecules
(Bover-Cid et al. 1999).
Certain drugs, for example, ephedrine and related compounds
(amphetamines, phentermine, mazindol, and fenfluramine) can
block the uptake of the biogenic amines NE and serotonin. Thus,
they are able to suppress appetite and food intake, and there is
interest in the development of thermogenic drugs that could act
on appropriate biogenic amines to control obesity safely without
increasing heart rate and blood pressure.

Quality Indicators
Biogenic amines are of interest and use to food scientists/food
technologists due to their use as food quality indicators. Aqueous

solutions of putrescine and cadaverine impart discernible and
objectionable odors at levels of 22 ppm and 190 ppm, respectively. The relative contribution of flavor from biogenic amines
to the overall food quality is not well established; rather, the
main focus has been on their possible use as chemical indicators of food quality. The relationship between the amounts of
particular biogenic amines in a food product and the extent of
food spoilage has been used to estimate a parameter known as
the chemical quality index (CQI) or the biogenic amines index
(BAI) for fish, and this takes into account the concentrations of
putrescine, cadaverine, histamine, spermine, and spermidine in
the fish sample (Mietz and Karmas 1977). On the basis of this,
the CQI is calculated as
Histamine + Putrescine +
Cadaverine
Chemical Quality Index [CQI] =
1 + Spermine + Spermidine
and a CQI value below 1 denotes a good quality product, a value
between 1 and 10 is considered mediocre, while a value ≥10
connotes spoilage. The CQI or BAI concept was extended further
as the beer BAI by taking into consideration the concentrations
of other biogenic amines, tyramine, β-phenylethylamine, and


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agmatine that occur in products like beer (Loret et al. 2005).
The beer BAI is calculated as
Cadaverine + Histamine + Tyramine +
Putrescine + Phenylethylamine + Trytamine
Beer BAI =
1 + Agmatine
Hitherto, a beer BAI value below 1 indicates a good quality
product, a value between 1 and 10 is considered as mediocre
quality, while a value ≥10 connotes spoilage. For meats, the
total content of putrescine, cadaverine, histamine, and tyramine
may be used to determine freshness, with levels ≤5.0 µg/g meat
indicating high quality products (Hernandez-Jover et al. 1997).

ANALYSIS METHODS
There are several analytical methods available for the detection, isolation, and quantification of biogenic amine levels in
food products. These include chromatographic, fluorometric,
and enzyme-linked immunosorbent assay (ELISA) methods.

Extraction and Chromatography
Various chromatographic methods have been used to analyze
biogenic amines. These methods all entail an initial extraction
step to recover the amines from the food product using organic

solvents, together with perchloric and trichloroacetic acids. The
extract thus obtained may then be subjected to chromatographic
separation methods such as ion-exchange column chromatography, paper chromatography, thin layer chromatography (TLC),
gas liquid chromatography, and high-pressure liquid chromatography (HPLC; Moret and Conte 1996, Cinquina et al. 2004, Paleologos and Kontominas 2004). TLC is considered as the simplest
of the chromatographic methods for the separation and quantification of biogenic amines (Shalaby 1996). It is effective, accurate, fairly rapid, relatively inexpensive, and can be applied to
large sample sizes. HPLC methods are commonly used method
for biogenic amine analysis because they are highly sensitive
and reproducible, although they require relatively more sophisticated and expensive technology, and are more time-consuming
and tedious to carry out compared with TLC (Shakila et al. 2001,
Lange et al. 2002).

Fluorometric Methods
Biogenic amines may also be determined by fluorometric methods. These methods also involve extraction, purification, and
separation of the compounds as in the chromatographic procedures. However, in order to quantify particular biogenic amines,
the eluant is mixed with a molecule to form a fluorescent product,
and the degree to which it fluoresces at particular wavelengths is
determined and used as a measure of the amount of the biogenic
amine in a food product by comparing with known standards
(Du et al. 2002).

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Biosensors
This technology is based on the recognition of biogenic amines
by specific ligands such as enzymes, antibodies, receptors, or
microorganisms. The binding of the amines to these ligands is
“sensed” by electrochemical, mass, optical, or thermal sensors,
and the degree of response is measured and correlated with the
amount of the substrate (amines) present in the raw material or
sample (Tombelli and Mascini 1998, Sarkadi 2009). Commonly

used biosensors for biogenic amines include putrescine oxidase
and DAOs (Carelli et al. 2007). These enzymes utilize O2 to react
with the amines and produce H2 O2 . The change in concentration
(of O2 or H2 O2 ) is typically measured as changes in electric
signal by electrodes (Prodromidis and Karayannis 2002).

Enzyme-Linked Immunosorbent Assay
Biogenic amines in foods may also be measured by the ELISA.
This method is based on the detection of N-amino derivatives of
histamine using antibodies. The N-amino derivatives are synthesized from histamine before analysis using compounds such as pbenzoquinone or propionic acid esters. There are presently commercial ELISA test methods that are available for the analysis
of histamine levels in wine, cheese, and fish (Serrar et al. 1995,
Aygun et al. 1999, Rupasinghe and Clegg 2007). The ELISA
method used to analyze wine sometimes yields slightly higher
results than the HPLC method; however, the ELISA method has
the advantage of being more rapid and less tedious to perform
than the HPLC methods.

EFFECTS OF FOOD PROCESSING
AND STORAGE
Food processing is an essential mechanism for controlling the
biogenic amine levels in foods. Because of their high thermal stability, once amines are formed, their concentrations will not be
significantly decreased during cooking and heating processes.
Processing techniques such as evisceration, postharvest handling, freezing, salting, and smoking can greatly affect the biogenic amine content and quality of the final product.

Evisceration
Evisceration can help control fish quality. For example, Atlantic
croacker and trout eviscerated directly after capture had a longer
shelf life than their iced un-eviscerated counterparts, and the
cadaverine and histamine concentrations were significantly less
in the eviscerated samples versus those of un-gutted fish samples

(Paleologos et al. 2004). This observation is due to a decrease in
the load of microorganisms associated with the eviscerated fish
compared with the un-eviscerated ones.

Temperature Control
Biogenic amine levels in food products are also greatly influenced by storage temperature. Postharvest handling techniques
like immediate icing are important in controlling biogenic amine


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levels in fish because at the reduced temperatures, both enzymatic and microbial activities are considerably reduced, so the
rates of formation of the biogenic amines and their accumulation
in the products are correspondingly reduced.


Salting
Salting can effectively inhibit biogenic amine formation particularly at high levels. In general, a higher salt content results in a
reduced biogenic amine formation. This is because the high salt
lowers the Aw of the milieu, and this can inhibit both the microbial and enzymatic activities that are required for the biogenic
amine formation.

Smoking
Smoking is a traditional method used to preserve fish. The fish
is smoked, then frozen, and transported. Histamine content is
found to increase during the smoking process and continues
to accumulate during subsequent freezing (Zotos et al. 1995).
Even though smoking may induce histamine production and
nitrosamines formation, the high temperature applied in smoking
can also curtail the growth and proliferation of harmful and/or
spoilage microorganisms (Martuscelli et al. 2009).

REGULATIONS
Since high level of biogenic amines, for example, histamine and
tyramine, are associated with food poisoning, their intake has to
be limited. Even though not all amines are equally toxic, and toxicological levels are hard to ascertain due to synergetic effects between amines, permissible limits have still been proposed. Thus,
fermented products that are prepared using good manufacturing
practices may be expected to contain histamine, tyramine, and
β-phenylethylamine concentrations of 50–100 , 100–800, and
30 mg/kg, respectively, and still considered safe and acceptable for human consumption (Nout 1994). Levels of histamine,
cadaverine, putrescine, tyramine, and β-phenylethylamine in
sauerkraut must not exceed 10, 25, 50, 20, and 5 mg/kg, respectively, and the total amount of biogenic amines in fish, cheese,
and sauerkraut must be less than 300 mg/kg (Shalaby 1996).
Furthermore, biogenic amine intake greater than 40 mg in a
meal is potentially toxic.

As biogenic amines are not extremely toxic or associated with
many fatal incidences, there are few general regulations controlling their concentration. Histamine, however, is a common
concern and different countries have set up their own standards
for these compounds in foods. For example, a level of 10 mg
histamine for every liter of wine is considered as acceptable in
Switzerland. In the United States, 20 mg histamine per 100 g
canned fish is considered unsafe for human consumption, and
50 mg histamine per 100 g canned fish is considered as a health
hazard by the Food and Drugs Administration (FDA 2001); the
European Economic Community has set the acceptable level of
histamine content in fish as 10–20 mg/100 g (Shalaby 1996);
the Canadian Food Inspection Agency, has set the action level

of histamine as 20 mg/100 g in fermented products and 10 mg/
100 g in scombroid fish products (CFIA 2005, Moret et al. 2005).

CONCLUSION
Biogenic amines are important food components found in many
fermented and non-fermented food production. They are present
as by-products of microbial activity in foods. The microorganisms are either present naturally, added for fermentation purposes, or are introduced through contamination. It is desirable
to minimize the formation of biogenic amines due to their adverse effects on human health and their contribution to food
spoilage and food losses. In particular, the levels of tyramine
and histamine need to be below a certain threshold level in order
to prevent toxic responses.
The study of biogenic amines is an ongoing field of research,
with several foci. Analytical methods are constantly being developed and/or improved in order to quantify their presence
more rapidly and accurately. Further studies are needed to generate the basic knowledge on the effects of ingestion, toxic limits, and interactions with other biological molecules, to facilitate the rationalization and formulation of more useful recommendations and regulations regarding their safe levels in food
products.

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