6A Amino Acids and Biogenic Amines 173
wine (Moreno-Arribas and Lonvaud-Funel 1999; Moreno-Arribas et al. 2000, 2003;
Constantini et al. 2006; Landete et al. 2007b).
CommercialO. oenistrains are selected for their oenological parameters, includ-
ing the absence of amino acid decarboxylases. According to the in vitro studies done
by Moreno-Arribas et al. (2003), none of the four commercial malolactic starter
cultures tested could produce histamine, tyramine or putrescine. Mart ́ın-Alvarez ́
et al. (2006) also compared inoculation with spontaneous malolactic fermentation
in 224 samples of Spanish red wine. They found that inoculation with a commer-
cial starter culture of lactic acid bacteriacould reduce the incidence of biogenic
amines compared to spontaneous malolactic fermentation in wines. Starter cultures
could eliminate indigenous bacteria, or could possibly degrade the biogenic amines
produced by the undesirable strains.
Other authors reported that no biogenic amine producers were present among
naturally occurringO. oenistrains isolated from wine and must (Moreno-Arribas
et al. 2003). Constantini et al. (2006) found noO. oeniamong 92 strains able to
produce biogenic amines in a broth medium. PCR screening was used by these
authors to confirm the absence of the respective decarboxylase genes.
In contrast, other authors found thatO. oeniis able to significantly contribute to
the overall histamine content of wines and also that the ability of this species to pro-
duce biogenic amines varies among strains (Coton et al. 1998; Guerrini et al. 2002).
Marcobal et al. (2004) isolated and identified a strain of theO. oenispecies, a pro-
ducer of putrescine and the capacity of another 42 strains of this species to produce
this amine has also been studied at a molecular level and the gene that encodes
biosynthesis of this amine was not present in any of them (Marcobal et al. 2004).
These results indicate that although this species may be involved in the production
of putrescine in wines, this is not a common property.
Amino acid decarboxylases are the enzymes responsible for the synthesis of
biogenic amines. These proteins and the genes that encode them have been studied
in several lactic acid bacteria in wine and are briefly discussed below.
Histidine decarboxylase (EC.4.1.1.22)is the enzyme that decarboxiles histi-
dine to produce histamine. The first decarboxylase enzyme of a wine lactic acid
bacterium was isolated from a histamine-producing strain by Lonvaud-Funel and
Joyeux (1994). Coton et al. (1998) purified this enzyme to homogeneity and pro-
vided molecular data. The gene sequence aided researchers to develop rapid and
specific detection systems based on polymerase chain reaction (PCR) to detect
potential histamine-producing bacteria from wine (Le Jeune et al. 1995; Coton
et al. 1998). Lucas et al. (2005) related histidine decarboxylase activity ofLacto-
bacillus hilgardii0006 isolated from wine, to the presence of an 80-kb plasmid
on which the decarboxylase gene was located as part of a four-gene cluster. These
authors suggest that a plasmid-encoded histidine decarboxylase could be transferred
horizontally, and that the location of the gene on an unstable plasmid may explain
the random distribution of histidine decarboxylase bacteria.
Tyrosine decarboxylase (EC.4.1.1.25), responsible for the production of tyramine
from tyrosine, was first investigated in a wine lactic acid bacteria by