6A Amino Acids and Biogenic Amines 171
Kanny et al. (2001) report that a normal individual can tolerate 129 mg/L of
histamine taken orally before symptoms occur, but only 7 g administered intra-
venously. However, there are studies that conclude that oral ingestion of biogenic
amines and wine tolerance are not related (Kanny et al. 2001; Jansen et al. 2003).
Rather, these authors propose that wine may contain compounds (ethanol and
acetaldehyde) that stimulate the release of histamine within the body. Amines are
also important in wine from a financial point of view, since they could cause
problems in commercial transactions (in their import and export). Although the
legal limits have not yet been established for any biogenic amine, some countries
have drawn up their own recommendations, especially for histamine. The exporta-
tion of wines to these counties could be paralyzed in the future, making biogenic
amines a potential economic threat. In fact, Switzerland and Austria reject wines
which contain more than 10 mg/L, and lower top limits have been recommended in
Germany (2 mg/L), Holland (3 mg/L), Belgium (5–6 mg/L) and France (8 mg/L)
(Lehtonen 1996).
Most wine-producing countries in the world, including Greece, Spain, Hungary,
Oregon (EEUU), Canada, South Africa, Italy and France, have investigated the
presence and concentration of biogenic amines in their wines (Gl `oria et al. 1998;
Soufleros et al. 1998; Mafra et al. 1999; Gerbaux and Monamy 2000; Haj ́os et al.
2000; Herbert et al. 2005; Bover-Cid et al. 2006; mo Dugo et al. 2006; Soufleros
et al. 2007; Smit 2007). In general, red wines have higher concentrations of biogenic
amines than white wines. In the former, these higher values are attributed to the
presence of lactic acid bacteria and malolactic fermentation. White wines, in gen-
eral, contain lower concentrations of amino acids and lower pH, due to the absence
of maceration during alcoholic fermentation. Evidently, other types of wines and
winemaking processes, such as Cava wines and other sparkling wines made by the
traditional method, and also biologically aged wines, are susceptible to the prob-
lem of biogenic amine formation, although there are very few studies of this in the
literature (Bodmer et al. 1999; Moreno-Arribas and Polo 2008).
6A.3.2 Microorganisms Associated with Biogenic Amine
Production in Winemaking
Potentially, all the microbial groups that participate in the winemaking process may
be associated with biogenic amine production. However, there is general agreement
that yeasts make a less significant contribution than lactic acid bacteria to the final
concentration of biogenic amines in wine.It is also clear that yeasts form different
biogenic amines to lactic acid bacteria. On the other hand, to date, there is much less
information about the biochemistry, genetics and regulation of amine production by
wine yeast, especially compared with the data available for lactic acid bacteria. The
formation of biogenic amines by wine microorganisms is discussed below.