172 M.V. Moreno-Arribas and M.C. Polo
6A.3.2.1 Yeast
A large variety of indigenous yeast species can grow and perform alcoholic fermen-
tation in wine, along with commercialSaccharomyces cerevisiaestrains. Few stud-
ies have been conducted on the formation of biogenic amines by yeasts, and most of
these only compared different yeast species and only quantified histamine (Torrea
and Anc ́ın 2002). Somavilla et al. (1986), using six yeast strains, demonstrated that
small amounts of histamine are produced during alcoholic fermentation and that the
association of yeasts and lactic acid bacteria can reduce the histamine level. The
highest histamine values (from 3.7 to 8.3 mg/L) were obtained when histidine was
added to the must (34 mg/L), in the other experiments histamine values were lower
than 1.2 mg/L. Vidal-Carou et al. (1990a) did not detect histamine formation either
during alcoholic fermentation although they did detect tyramine formation, but at
very low levels (0.60 mg/L). In contrast, other authors disagree with the hypothesis
that biogenic amines are formed by lacticacid bacteria during malolactic fermenta-
tion (Buteau et al. 1984). Torrea-Go ̃ni and Anc ́ın-Azpilicueta (2001) found a slight
biogenic amine production bySaccharomyces cerevisiaedepending on the strain.
Recently, Landete et al. (2007b) screened 36 strains of yeast isolated from must
and wines to produce biogenic amines. The yeast tested included strains belong-
ing to the generaAureobasidum, Candida,Hanseniaspora, Hansenula, Kloeck-
era,Metschnikowia, Pichia,Rhodotorulaand strains of the speciesSaccharomyces
cerevisiae, S. cerevisiaevar.bayanus, S. cerevisiaevar.chevalieri and S. cerevisiae
var.steiner. No biogenic amines were produced by any of the yeast strains tested
under the conditions present in this screening test, in synthetic medium, grape
must or wine. These results are consistent with previous studies in which neither
histamine, tyramine or putrescine production were detected in 50 yeast strains
isolated from grapes and/or wine, includingSaccharomyces cerevisiaeand other
non-Saccharomycesyeasts, while some of these yeasts varied in their capacity to
form phenylethylamine (Caruso et al. 2002; Granchi et al. 2005). These findings,
therefore, indicate that yeast does not appear to be at least the direct origin of most
amines found in wines.
6A.3.2.2 Lactic Acid Bacteria
Extensive research has been done to correlate biogenic amine production in wine
with species of lactic acid bacteria involved in the winemaking process. The first
studies on histamine production by lactic acid bacteria indicate thatPediococ-
cus were the main producers (Farias etal. 1993). Recent studies by Landete
et al. (2007b) are in accordance since, although the percentage ofPediococcus
species capable of producing histamine seems to be low, some strains are responsible
for the highest concentrations. However, today it is well known thatLactobacillus,
LeuconostocandOenococcusspecies are also implicated in biogenic amine pro-
duction in wine. Different strains ofLactobacillus hilgardii, L. brevis. L. buchneri
andL. malihave been found to be able to produce a variety of biogenic amines in