Food Biochemistry and Food Processing (2 edition)

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43 Biogenic Amines in Foods 825

problems, palpitations, headaches, rashes, and blood pressure
problems (Bodemer et al. 1999, Du Toit 2005). In the process,
fermentation microorganisms such asSaccharomyces cerevisiae
andS. ellipsoideushydrolyze carbohydrates into simpler sugars
that are further degraded into ethanol and a variety of flavor com-
pounds. The beverages are then stored and aged for various time
periods. In wines, the most common biogenic amines are his-
tamine, tyramine, and putrescine, which are formed by bacterial
strains such asPediococcus spp., L. brevis, L. buchneri, Leu-
conostoc mesenteroides, andOenococcus oeni(Guerrini et al.
2002, Moreno-Arribas et al. 2003).
The production of beer entails an initial mashing step to pro-
duce the wort, which is subsequently inoculated with yeast (e.g.,
S. carlsbergensisorS. cerevisiae) to potentiate the fermenta-
tion. Histamine, tyramine, putrescine, and cadaverine have been
found in certain beers and because alcohol enhances the ab-
sorption of biogenic amines in humans, a legal upper limit for
biogenic amine content of 2 mg per alcoholic beverage has been
established as guide for manufacturers of the products (Moreno-
Arribas et al. 2003).
The formation and accumulation of biogenic amines in beer
is due to several factors, such as the quality of the raw material
and microbial contaminations. Thus, lactic acid bacteria form
putrescine, cadaverine, histamine, and tyramine during fermen-
tations; storage and aging may lead to further increases in the
levels of these compounds in the food products (Tabor et al.
1958, Bodemer et al. 1999).
The formation and/or accumulation of biogenic amines in
foods may be controlled by adopting proper good sanitary and
hygienic practices during food handling and processing, to con-
trol microbial contaminations and prevent temperature abuse
during storage and transportation.

Meat Products

Fermentation is also used as a traditional preservation method
to extend the shelf life of meat. Examples of fermented meats
include ham, salami, pepperoni, chorizo, thuringer, bologna, and
cervelat (Potter and Hotchkiss 1998). Sausages are classified as
semi-dry and dry sausages, based on fermentation techniques.
The semi-dry sausages are characterized by a relatively higher
water activity (Aw=0.90–0.94) and a relatively higher pH
(≤5.0), as opposed toAwof 0.85–0.91 and a pH of≤4.5 for
dry sausage. The semi-dry sausages are usually produced using
a lactic acid bacteria starter culture over a relatively shorter fer-
mentation period, while dry sausages are fermented over much
longer time periods using the endogenous microflora associated
with the raw material. The biogenic amines histamine, tyra-
mine, tryptamine, cadaverine, putrescine, 2-phenylethyamine,
spermidine, and spermine have all been found in dry sausages.
However, their levels and distribution vary depending on factors
such as quality of the raw material and the availability of pre-
cursor molecules to serve as substrates (Santos 1996, Suzzi and
Gardini 2003).
Raw meats tend to have lower amounts of biogenic amines
such as spermine and spermidine unlike fermented products, and
some industry-derived commercial products tend to have rela-

tively higher amounts of biogenic amines than their nonindustry-
produced traditional products (Lakritz et al. 1975). The mi-
croorganisms responsible for biogenic amines production in fer-
mented meats includeEnterobacteriaceae,Enterococci,Lacto-
bacilli, Pediococcus,andPseudomonads. The lactic acid bac-
teria strain,LactobacilliandEnterococci,are more associated
with tyramine production, whileEnterobacteriaceaeis more
connected with high putrescine and cadaverine levels in fer-
mented meat products (Suzzi and Gardini 2003).
It is possible to have the microbial load associated with fer-
mented meat products to be low; however, the enzymes (pro-
teases and decarboxylases) released by microorganisms into the
products may survive the fermentation process and cause con-
tinued production of biogenic amines in the finished products.
Storage of the raw materials intended for fermentation at low
temperatures is a good practice to control both microbial pro-
liferation and enzymatic activity that lead to biogenic amines
formation in the products. Fermented meat products made from
raw material that have been exposed to high temperatures tend
to have significantly high contents of biogenic amines such as
tyramine, cadaverine, and putrescine.
In general, high temperature, high pH, and low salt content
can enhance biogenic amine formation. Higher temperatures
favor proteolytic, lipolytic, and decarboxylation reactions. For
instance, the amount of biogenic amine content is higher in prod-
ucts prepared at 15◦C compared with similar products made at
4 ◦C, and more histamine is formed during ripening at 18◦Cas
compared with processing at 7◦C; biogenic amine formation
is also less at lower pHs, because the decarboxylation of his-
tidine tends to be relatively low at acidic pH (Maijala et al.
1995). Thus, amine-negative lactic acid bacteria are preferred
as starter culture to rapidly produce acids to decrease the pH in
the product(s) and suppress biogenic amine formation (Battcock
and Azam-Ali 1998). A high salt content also generally inhibits
microflora growth because the reduced water activity makes
the product less conducive for the growth and proliferation
of microbes; consequently, low levels of biogenic amine are
formed.
Other factors may also contribute to the biogenic amine ac-
cumulation. For example, bacteria growth and metabolism are
enhanced by the presence of glucose, while the addition of sul-
phites or nitrites suppresses the formation of cadaverine, pu-
trescine, and cadaverine. In the case of sausages, the diameter
of the product may affect biogenic amine formation, because it
impacts the salt content and water activity of the product. Thus,
sausages with bigger diameters tend to have greater tendency for
biogenic amine production and accumulation than smaller-sized
sausages (Battcock and Azam-Ali 1998).
In addition to meats, fish are also fermented into products
such as faseekh, fish paste (e.g., bagoong, patis, and shrimp
paste), and fish sauce (e.g., joetgal, shottsuru, garum, nuoc mam,
and nam pla). Like their meat counterparts, these products also
may contain biogenic amines. Histamine, putrescine, cadaver-
ine, tyramine, spermidine, and spermine have all been detected
in fermented fish products and their levels increased with stor-
age (Mah et al. 2002). However, the amounts of biogenic amines
found in these products (0–70 mg/kg sample) are considered as
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