368 M. Ugliano and P.A. Henschke
Ta b l e 8 D. 1 0Principal characteristics of selected non-SaccharomycesyeastsGenus species Characteristics
Torulaspora delbrueckii (Candida colliculosa) Osmotolerant, low volatile acidity
Kluyveromyces thermotolerans Bioacidification, aroma enhancement
Candida stellata High glycerol, preferential fermentation
of fructose
Issatchenkia orientalis (Candida krusei) Malic acid degradation,
aroma enhancement
Metschnikowia pulcherrima (Candida pulcherrima) Aroma enhancement
Hanseniaspora uvarum (Kloeckera apiculata) Flavour complexity
(potential to increase volatile acidity)
Schizosaccharomyces malidevorans Deacidification (malic acid)
(Schizosaccharomyces pombe)
Dekkera bruxellensis (Brettanomyces bruxellensis) Flavour complexity
Pichia anomala, Pichia fermentans, Pichia Aroma enhancement
membranifaciens,Issatchenkia terricola,Pichia
subpelliculosa, Williopsis staurnus
Jolly et al. 2003b; Moreno et al. 1991).Torulaspora delbrueckiihas been isolated
from high sugar musts and this osmotolerant property is exploited in the fermenta-
tion of botrotized wines (Henschke and Dixon 1990; Lafon-Lafourcade et al. 1981;
Bely et al. 2008). Under hyperosmotic stress conditionsTorulaspora delbrueckii
typically retains the ability to produce high glycerol and low acetic acid when com-
pared toSaccharomyces, although cofermentation withSaccharomycesis necessary
to yield high ethanol concentrations.Torulaspora delbrueckiiis a component of a
mixed yeast culture withSaccharomyces cerevisiaeandKluyveromces thermotol-
eransthat has been commercialized by Chr. Hansen (Viniflora©R HARMONY.nsac
and MELODY.nsac).
Kluyveromces thermotoleransis an acid-producing yeast, which has a mod-
erate to high tolerance to ethanol (<13.5 vol.%) when compared to other non-
Saccharomycesspecies. Strains of this species can produce up to 7 g/L of lactic
acid, and has the potential to balance low acidity musts. Strains ofKluyveromces
thermotoleransare generally characterised by moderate ester, higher alcohol and
acetic acid production and low acetaldehydeand off-flavour production, suggesting
suitability to wine production (Ciani et al. 2006; Kapsopoulou et al. 2007; Mora
et al. 1990; Zeeman et al. 1982).Kluyveromyces thermotoleransis a component of a
mixed yeast culture withSaccharomyces cerevisiaecommercialized by Chr. Hansen
(Viniflora©RSYMPHONY.nsac).
Candida stellatais widely distributed in musts world-wide, and while it can con-
sume 2-ketogluconate, a sulfite binding substance produce in grapes and musts by
Botrytis cinereaand acetic acid bacteria, some strains are tolerant to sulfite and
can produce large concentrations of 2-methyl-1-propanol, rendering such strains
of this yeast undesirable in fermentation (Holloway et al. 1992;). However, more
recent studies on other isolates have revealed several potentially useful winemaking
characteristics, including high glycerol and succinic acid production, a small reduc-
tion in final ethanol, preference for fructose consumption fermentation, moderate