Food Chemis y: X 9 (2021) 100116
A ailable online 9 Feb ua y 2021
2590-1575/© 2021 The Au ho (s). Published by Else ie L d. This is an open access a icle unde he CC BY license (h p://c ea i ecommons.o g/licenses/by/4.0/).
The e ec s o Saccha omyces ce e isiae s ains ca ying alcoholic
e men a ion on he e men a i e and a ie al a oma p o iles o young and
aged Temp anillo wines
Ma ie Dena
a
, Dolo es P´
e ez
b
,
c
,
d
, Jos´
e Ma ía He as
b
, Ampa o Que ol
c
, Vicen e Fe ei a
a
,
*
a
Labo a o y o A oma Analysis and Enology (LAAE), Depa men o Analy ical Chemis y, Uni e sidad de Za agoza, Ins i u o Ag oalimen a io de A ag´
on (IA2)
(UNIZAR-CITA), c/Ped o Ce buna 12, 50009 Za agoza, Spain
b
Lallemand Bio S.L., 08028 Ba celona, Spain
c
Ins i u e o Ag ochemis y and Food Technology (IATA-CSIC), 46980 Pa e na, Valencia, Spain
d
Es aci´
on Expe imen al Ag opecua ia Mendoza (EEA), Ins i u o Nacional de Tecnología Ag opecua ia (INTA), 5507 Luj´
an de Cuyo, Mendoza, A gen ina
ARTICLE INFO
Keywo ds:
Fe men a ion
Longe i y
F ui y es e s
Vola ile phenols
No isop enoids
Vanillin
S ecke aldehydes
E hyl leuca e
ABSTRACT
Ten di e en Saccha omyces ce e isiae s ains e men ed semi-syn he ic mus s con aining a Polyphenolic and
A oma P ecu so F ac ion (PAF) ex ac ed om Temp anillo g apes. A oma compounds we e s udied by Gas
Ch oma og aphy (GC), GC-Ol ac ome y and GC-Mass Spec ome y (MS), du ing e men a ion by apping
ola ilized a oma, immedia ely a e e men a ion and a e accele a ed aging. Vola iles los by e apo a ion
du ing e men a ion a e mos ly e men a i e compounds and no g ape- ela ed odo an s. Isobu anal and some
es e s a e mos ly los du ing e men a ion. In many cases he impac o yeas s ain is e iden only a e aging.
S ains could be classi ied in o 3 majo clus e s wi h ma ked di e ences in e men a i e and a ie al p o iles.
Linalool and ge aniol we e ound o ha e e men a i e o igin. S. ce e isiae yeas s ains can e ec i ely modula e
a ie al a oma, likely h ough speci ic enzyma ic ac i i ies ac ing on g ape phenolic acids and no isop enoid
a oma p ecu so s and may be speci ically used o mi iga e some aging- ela ed o odou s, such as massoia
lac one, guaiacol o TDN.
1. In oduc ion
Wine a oma is i s mos ou s anding senso y p ope y and is essen ial
o i s quali y and di e en ia ion (Cha e s & Pe ig ew, 2007).
Al hough he numbe o ola ile molecules which can be a pa o he
ola ile ac ion o wines is e y la ge, exceeding mos likely se e al
housands, i has been sugges ed ha 70 di e en odou chemicals a e
hose playing majo oles on he a oma ic p ope ies o wines (De-la-
Fuen e-Blanco, S´
aenz-Na ajas, & Fe ei a, 2020).
Quan i a i ely, he mos abundan wine odo an s a e alcoholic
e men a ion (AF) by-p oduc s, pa icula ly highe alcohols, e hyl es e s
and ace a es, some ca bonyls and acids. By con as , many o he ele-
an odo an s de i e om g ape speci ic p ecu so s and can be p esen
a e y limi ed concen a ions, wi hin he ng/L ange in he case o
poly unc ional me cap ans, ew hund eds o ng/L in he case o β-ion-
one, a ound he ew
μ
g/L in he case o β-damascenone o below 0.2 mg/
L in he cases o e penols, ola ile phenols and anillin de i a i es
(Fe ei a & Lopez, 2019; Ruiz e al., 2019).
The e a e also ele an di e ences be ween compounds ega ding
he ime a which hey a e o med o libe a ed. Mos e men a ion by-
p oduc s de i e om yeas amino acid and a y acid me abolisms
(Swiege s, Ba owsky, Henschke, & P e o ius, 2005), so ha hey a e
signi ican ly o med om he ea ly s ages o AF and a e known o be
pa ly los by e apo a ion (G´
omez-Plaza, Ma ínez-Cu illas, & Laencina,
1993; Gue ini e al., 2016). Some o he compounds, such as he mon-
o e penoids ge aniol and linalool, a e di ec ly libe a ed om
g ape-speci ic glycosyla ed p ecu so s du ing e men a ion by he ac ion
o yeas β-glucosidases (Guna a, Bi eu , B illoue , Bayono e, & Co -
donnie , 1988). Poly unc ional me cap ans a e also byp oduc s o AF
and may be also pa ially los by e apo a ion du ing e men a ion.
On he con a y, he o ma ion and accumula ion o some o he
ola iles equi es aging ime. In ac , a quan i a i ely ele an g oup o
a oma compounds and a oma p ecu so s a e subjec ed o se e al slow
chemical eac ions such as acid hyd olysis, es e i ica ion o in a-
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (V. Fe ei a).
Con en s lis s a ailable a ScienceDi ec
Food Chemis y: X
jou nal homepage: www.sciencedi ec .com/jou nal/ ood-chemis y-x
h ps://doi.o g/10.1016/j. ochx.2021.100116
Recei ed 20 Ma ch 2020; Recei ed in e ised o m 24 Decembe 2020; Accep ed 19 Janua y 2021
Food Chemis y: X 9 (2021) 100116
2
molecula ea angemen s ha will g ea ly a ec wine a oma p o ile.
This is he case o mos no isop enoids and anillin de i a i es, such as
β-damascenone and TDN o anillin and ace o anillone, whose le els
inc ease du ing aging as he consequence o di e en ans o ma ions o
g ape ca o enoid me aboli es (Win e hal e & G¨
ok, 2013) o o g ape
glycosyla ed p ecu so s (Fe ei a & Lopez, 2019). No ably, i has been
ecen ly obse ed ha he yeas gene a plays a majo ole in he mod-
ula ion o TDN concen a ion in wine a e some ime o aging (Oli ei a
& Fe ei a, 2019). A hi d g oup o odo an s su e ing changes du ing
aging a e labile molecules, such as linalool o ge aniol, which deg ade
du ing aging o o m
α
- e pineol, ne ol o 1,8-cineole (Wa e house,
Sacks, & Je e y, 2016). O he g oups o odo an s deeply a ec ed by
aging a e ui y es e s and ace a es. The ace a es o highe alcohols a e
quickly hyd olysed and hei le els soon ade away. On he con a y, he
e hyl es e s o b anched acids - isobu y ic, 2-me hylbu y ic and iso-
ale ic acids - and o o he mino acids, slowly and s eadily inc ease by
es e i ica ion o hei p ecu so acids wi h e hanol (Díaz-Ma o o,
Schneide , & Baumes, 2005). The exis ence o all hese p ocesses makes
ha aging ime should be conside ed as an impo an ac o o assess he
ole o yeas s on wine a oma modula ion.
Saccha omyces ce e isiae is he mic o-o ganism mos widely s udied
ega ding i s senso y impac on wine (Temp`
e e e al., 2018). Howe e ,
mos p e ious s udies ha e deal wi h he sho - e m impac o his yeas
on bo h e men a i e and a ie al a oma p o iles (Game o, He n´
andez-
O e, Que ol, & Fe ei a, 2011; Gammacu a, Ma chand, Moine, & de
Re el, 2017; Molina e al., 2009), neglec ing aging e ec s and also
possible di e ences in he a oma p o iles los by e apo a ion, which
could be pa icula ly ele an in lab-scale e men a ions. Bo h aspec s
will be specially add essed in he p esen wo k whose objec i e is o
e alua e he impac o S. ce e isiae yeas s ains on he e men a i e and
a ie al a oma o Temp anillo wine du ing and a e e men a ion, and
a e a pe iod o accele a ed aging.
2. Ma e ial and me hods
2.1. Reagen s and s anda ds
Dichlo ome hane and me hanol (≥99%) Dis o-Pes icide esidue
g ade we e supplied by Me ck (Da ms ad , Ge many). ACS quali y ab-
solu e e hanol was pu chased om Pan eac (Ba celona, Spain). 2-
Bu anol (≥99%), 4-me hyl-2-pen anol (99%), 4-hyd oxy-4-me hyl-2-
pen anone (99%), e hyl hep anoa e (99%) and hep anoic acid (99%)
we e used as in e nal s anda ds o majo compounds analysis. 2-Oc a-
nol (99.5%), 3-oc anone (99%) and 3,4-dime hylphenol (99%) we e
used as in e nal s anda ds o mino compounds analysis. They we e
pu chased om Me ck as well as he chemical s anda ds used in his
s udy (>98%), excep o TDN which was syn hesized by Synchem UG
&Co (Felsbe g, Ge many) wi h a pu i y o 80%. LiCh olu EN esin
ca idges we e ob ained om Me ck (Da ms ad , Ge many). Sep Pak-
C18 esins, p epacked in 10 g ca idges we e om Wa e s (I eland).
2.2. Semi-syn he ic mus p epa a ion
2.2.1. P ecu so s and phenols ex ac ion om Temp anillo g apes
Phenolic and A oma p ecu so F ac ion (PAF) was ex ac ed om
Temp anillo g apes mis elle p epa ed a he ICVV o Log o˜
no (Spain)
ollowing he p ocedu e de eloped by Aleg e e al. (2020).
2.2.2. Syn he ic g ape mus
Syn he ic mus was p epa ed as desc ibed in He n´
andez-O e, Bely,
Cacho, and Fe ei a (2006). Ni ogen con en was adjus ed by mixing
220 mg/L o (NH
4
)
2
HPO
4
, and a mix u e o amino acids esembling he
a e age p o ile o Temp anillo g ape a ie y (He n´
andez-O e, Cacho, &
Fe ei a, 2002). Syn he ic glu a hionyla ed and cys einila ed (Cys)
p ecu so s o 3-me cap ohexanol (MH) and 4-me cap open an-2-one
(MMP) we e added om a solu ion in MilliQ wa e , (0.1 mg/L Cys-
MH, 0.05 mg/L Cys-MMP, 1 mg/L Glu-MH, 0.05 mg/L Glu-MMP).
A e pH adjus men o 3.5 wi h NaOH, syn he ic mus was s e ilized
by il a ion (0.45
μ
m) inside a e ical lamina low chambe . PAF was
dealcoholized, esuspended in s e ile dis illed wa e and added o syn-
he ic mus a 10% ( / ).
2.3. Wine elabo a ion
2.3.1. Yeas s s ains, p e-cul u e condi ions and yeas g ow h moni o ing
Ten S. ce e isiae s ains (Lallemand Bio SL, Mad id, Spain) we e used:
Lal in ICV D254™ (D254), Lal in Clos™ (CLOS), U a e m HPS™ (HPS),
Eno e m BDX™ (BDX), Lal in Rhˆ
one2056® (RHONE), Lal in ICV D80™
(D80), Lal in 71B™ (71B), Lal in Pe sy™ (PERSY), Lal in ICV OKAY™
(OKAY), IONYS w ™ (IONYS). Ac i e d y yeas s we e ehyd a ed in
dis illed wa e a 37 ◦C o 20 min unde agi a ion, 100
μ
L o hese
cul u es we e pla ed on Glucose Pep one Yeas (GPY) and incuba ed a
25 ◦C o 24 h. P e-cul u es we e p epa ed by inocula ing se e al col-
onies in o 5 mL o GPY b o h a 25 ◦C o 24 h. A e quick cen i uga-
ion, GPY b o h was disca ded and colonies we e esuspended in
dis illed wa e . Li ing cells we e quan i ied by low cy ome y and each
e men e o syn he ic mus was inocula ed a 10
6
li ing cells/mL. Cell
iabili y and i ali y we e moni o ed by low cy ome y (Tilloy, O iz-
Julien, & Dequin, 2014).
2.3.2. Expe imen al design
All he e men a ions we e ealized in iplica es. Fe men a ion wi h
CLOS and IONYS we e epea ed in mus wi hou PAF addi ion, whose
olume was eplaced by s e ile dis illed wa e . Un e men ed con ol o
syn he ic mus wi h and wi hou PAF was also included in duplica es.
Expe imen al p ocedu e is summa ized in Fig. S1.
2.3.3. Fe men a ion sys em
Fe men a ions we e ca ied ou in 100 mL glass e men e s con-
aining 50 mL o syn he ic mus so ha headspace ep esen ed 50% o
he o al olume. Fe men e s we e igh ly closed wi h a pe o a ed sili-
cone co k in which an ai lock (Mic omal a S.L., Mad id, Spain) was
inse ed. Needles and 5 mL sy inges we e inse ed in o he silicon co k o
allow sampling wi hou opening he e men e .
2.3.4. Fe men a ion moni o ing
Fe men a ions we e ca ied ou a 25 ◦C, unde agi a ion a 150 pm
using a magne ic s i e . CO
2
elease was moni o ed by weighing. The
main chemical e men a i e pa ame e s we e analysed a he beginning
and a he end o e men a ion, including suga s, acids and alcohols by
UHPLC as desc ibed in Su e al. (2019).
2.3.5. End o e men a ion and accele a ed aging
A he end o e men a ion, samples we e cen i uged and condi-
ioned o accele a ed anoxic aging in o a ee-O
2
chambe Jacomex
(Dagneux, F ance). Samples we e placed in o 18 mL glass ubes capped
wi h non-me allic sc ew caps and bagged in high densi y plas ic bags
con aining oxygen sca enge s Anae oGenTM (The mo Scien i ic, USA)
and incuba ed a 50 ◦C o 5 weeks. This me hodology was de eloped by
Vela, He n´
andez-O e, F anco-Luesma, & Fe ei a, (2017) and was
success ully applied o he aging o e men ed media (Oli ei a & Fe -
ei a, 2019). I was obse ed ha 5 weeks o anoxic aging a 50 ◦C we e
oughly equi alen o one yea o aging a oom empe a u e.
2.4. Analysis o he a oma e apo a ed du ing e men a ion
A p e-pu i ied s anda d SPE ca idge illed wi h 160 mg LiCh olu -
EN esin was lodged in o he ai lock o he e men e s o ap ola iles
emi ed du ing e men a ion. When his inished, he ca idge was
emo ed, d ied and elu ed wi h 1.6 mL o dichlo ome hane con aining
5% ( / ) o me hanol.
Fo GC-Ol ac ome y (GC-O), a single ex ac was p epa ed by
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
3
mixing 100-uL o each one o he o y ex ac s ob ained om he 40
di e en e men a ions. The ex ac was ca e ully concen a ed by
e apo a ion o he sol en unde ni ogen low un il a 0.2 mL inal
olume and injec ed in o he GC-O sys em, as desc ibed in San-Juan,
Pe ’ka, Cacho, Fe ei a, & Escude o, (2010). Ol ac ome ic sco es we e
ob ained by combining in ensi y and equency as desc ibed by D a -
nieks, (1985). The odo an s we e iden i ied by compa ison o hei de-
sc ip o s, ch oma og aphic e en ion index in DB-WAX and DB-5
columns and mass spec a wi h hose o pu e e e ence compounds.
Fo quan i ica ion o he odo an s p esen in he ex ac s, hese we e
spiked wi h he in e nal s anda ds, concen a ed by e apo a ion unde
ni ogen up o 0.2 mL and analyzed. Majo e men a i e compounds
we e di ec ly quan i ied by GC-FID analysis unde he condi ions
desc ibed in O ega, L´
opez, Cacho, & Fe ei a, (2001). Mino com-
pounds we e quan i ied by GC–MS analysis o he ex ac s in a Shimadzu
QP2010 (Quio o, Japan) equipped wi h a DB-WAXe GC-column om
Agilen (San a Cla a CA, USA), 30 m ×0.25 nm, 0.5
μ
m o ilm hickness,
p eceded by a 2 m ×0.25 mm uncoa ed p e-column. Ca ie gas was
Helium a 1.26 mL/min. Injec ion olume was 1
μ
L in spli mode ( a io
1/30). Injec o empe a u e was 250 ◦C. Ch oma og aphic o en em-
pe a u e was ini ially a 30 ◦C o 1 min, hen aised a 1 ◦C/min o
35 ◦C, held o 1 min, hen a 1 ◦C/min o 40 ◦C, a 15 ◦C/min o 55 ◦C,
held o 5 min, a 15 ◦C/min o 72 ◦C, held o 5 min, a 15 ◦C/min o
150 ◦C, held o 15 min. The Ion sou ce was kep a 220 ◦C and he
in e ace a 230 ◦C. The mass analyse was se in single ion moni o ing
mode. The comple e lis compounds quan i ied in bo h p ocedu es,
including he m/z a ios selec ed o MS quan i a ion is a ailable in
Table S1.
2.5. Analysis o majo and mino ola ile compounds
Majo me aboli es o alcoholic e men a ion (highe alcohols and
hei ace a es, ola ile a y acids and hei e hyl es e s, b anched a y
acids and hei e hyl es e s, ace oin, diace yl, and ace aldehyde), usually
p esen in wines a le els abo e 0.2 mg/L, we e analysed by he GC-FID
analysis o a dichlo ome hane mic oex ac as desc ibed by O ega,
L´
opez, Cacho, & Fe ei a, (2001).
Mino a oma compounds p esen in wine a le els 0.1–200
μ
g/L
(b anched e hyl es e s, e penes, no isop enoids, anillin de i a i es,
ola ile phenols) we e analysed by GC–MS analysis o a SPE ex ac
ollowing he p o ocol desc ibed by L´
opez, Azna , Cacho, & Fe ei a,
(2002). The comple e lis o quan i ied is a ailable in Table S1.
2.6. S a is ical analysis
S a is ical analysis and g aphics we e ealized using R so wa e
(RS udio Team, 2020). Signi icance was de e mined by analysis o
a iance (ANOVA) using ano a unc ion om ca package ( .3.0.2) and
Tuckey’s hones ly signi icance di e ence es was pe o med on ANOVA
esul s, using HSD. es unc ion om ag icolae package ( .1.3.1). P in-
cipal Componen Analysis (PCA) we e pe o med and plo ed using
ac oex a package ( .1.0.5).
3. Resul s
3.1. Odo an s los du ing e men a ion
In o de o assess he ype and amoun s o odo an s los du ing
e men a ion, a small ap was ins alled in he e men e s. A GC-O
sc eening p ocedu e was ca ied ou on an ex ac ob ained by mixing
small aliquo s o all he ex ac s ob ained in he expe imen . Resul s a e
summa ized in Table 1. O e all, wen y- wo odo an s we e de ec ed
wi h GC-O sco es abo e 20%. Ele en ou o he wen y- wo odo an s
we e iden i ied a maxima le el o con idence. In i e o he cases, some
o he iden i y c i e ia could no be comple ely ul illed because o
di e en easons, such as excessi ely low le els o ge a good MS
spec um (Z and E-2-nonenals), co-elu ion in he non-pola column
(c esol and guaiacol) o disc epancy in he odou (e hyl oc ano e). The
iden i ica ion o he six less in ense odo an s was based only on he
coincidence o he odou and RI o he odo an in he pola column wi h
hose o he s anda d, and should be conside ed en a i e.
The mos in ense odo an s we e mainly by-p oduc s o alcoholic
e men a ion: isoamyl alcohol and 2-phenyle hanol, 3-me hylbu anal,
and a nume ous g oup o es e s (isop opyl and isoamyl ace a es, and he
e hyl es e s o isobu y ic, bu y ic, hexanoic, oc anoic and decanoic
acids). Apa om hese, c esol and Z and E-2-nonenals we e also be-
ween he wel e mos in ense. The o igin o c esol is no clea , bu i
could be a b eakdown p oduc o he polyphenols p esen in e men a-
ion media; while nonenals a e known de i a i es o he au o-oxida ion
o g ape a y acids. Resul s, he e o e, con i m ha he mos ele an
odo an s pu ged ou du ing e men a ion a e by-p oduc s de i ed om
yeas me abolism o g ape a y acid au o-oxida ion and no a ie al
a oma compounds eleased om speci ic a oma p ecu so s in g ape,
such as e penols, no isop enoids o poly unc ional me cap ans.
3.2. Quan i a i e assessmen o he ola iles los du ing e men a ion
The amoun s o eigh een odo an s apped in he ca idges ins alled
in he PAF-con aining e men e s a e summa ized in Table 2. In gene al,
le els we e low and in some cases we e a ec ed by high imp ecision.
The o al mass o ola iles apped in he ca idges anged om a ound
1 mg (D80) o a ound 2 mg (71B). The majo ola ile was iso-
bu y aldehyde, which accoun s o mo e han 50% o he o al ola iles
apped. Fou o he odo an s, isop opyl ace a e, isoamyl ace a e, iso-
bu anol, and isoamyl alcohol can be also ound a le els abo e 100
μ
g.
The le els o some ola iles eleased we e signi ican ly ela ed o he
Table 1
Iden i ica ion o odo an s pu ged ou du ing e men a ions and apped in
LiCh olu -EN ca idges placed be o e he Mulle al es o he e men e s. The
GC-O expe imen was ca ied ou on an ex ac made by mixing he elua es o he
di e en aps. Re en ion indexes (RI) in DB-WAX and DB-5 columns, iden i i-
ca ions, ol ac ome ic sco es (MF%) and odo desc ip o s a e p esen ed in he
able. Compounds a e ma ked by le e s acco ding o he eliabili y o hei
iden i ica ion (see legend).
RI (DB-
WAX)
RI (DB-
5)
Compound* MF
(%)
Odo desc ip ion
1220 <900 isoamyl alcohol
A
87 cheese, ancid
940 <900 3-me hylbu anal
A
78 cheese, ancid
935 <900 isop opyl ace a e
A
62 ui y, s awbe y
975 <900 e hyl isobu y a e
A
62 ui y, s awbe y
2097 c esol
B
55 phenolic, lea he
1929 1115 2-phenyle hanol
A
53 lo al, ose
1039 <900 e hyl bu y a e
A
46 ui y, s awbe y
1241 988 e hyl hexanoa e
A
45 ui y
1513 1133 Z-2-nonenal
B
38 ancid, cucumbe
1128 <900 isoamyl ace a e
A
37 banana
1441 1193 e hyl oc anoa e
B
35 mush oom, plas ic,
humidi y
1544 1153 E-2-nonenal
B
29 a , ancid
1621 1391 e hyl decanoa e
A
29 soap
1696 <900 2 and 3-me hylbu y ic
acid
A
26 swea , ancid
1842 guaiacol
B
26 smoky, bu n
966 <900 e hyl p opanoa e
A
24 ui y
1425 1-nonen-3-one
C
24 mush oom,
unde g ow h
1468 decanal
C
24 g ass, lo al
1953 Z-whisky lac one
C
24 spicy
2169 2-phenoxye hanol
C
24 ancid, ca on
2245 4- inylguaiacol
C
24 spicy
2214 so olon
C
22 spicy
* A: iden i ica ion conclusi e. Expe imen al RIs in wo columns, odou and MS
co esponded o he one ob ained wi h he pu e chemical s anda d; B: iden i y
highly likely, one o he p e ious c i e ia ( wo RIs, odou , MS) ailed; C: en a i e
iden i ica ion based on RI on a single column, odou and p e ious li e a u e.
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
4
yeas s ain, al hough in mos cases di e ences we e no e y high.
Majo di e ences co espond o he s ain IONYS, whose ca idges
con ained highes le els o ace a es and o some e hyl es e s, ollowed by
he s ain 71B, wi h maxima le els o aldehydes.
As expec ed, ola iles pu ged du ing e men a ion a e mos ly non-
pola a oma compounds such as es e s, ace a es o ca bonyls. Only
wo acids and wo alcohols we e ound amongs he quan i iable ola ile
compounds. In hese cases, he amoun s e apo a ed co esponded o
e y small ac ions o he ola iles p oduced. In he pa icula case o
isoamyl alcohol, he 664
μ
g ound in he ca idge o OKAY s ain
co espond o 13 mg/L in he 50 mL o liquid. Conside ing ha he
ecen ly e men ed wine con ained a ound 190 mg/L o his compound
(Table 2, Supplemen a y ma e ial), i can be es ima ed ha less han 5%
o he o al amoun o isoamyl alcohol p oduced was e apo a ed. On he
opposi e side, isobu y aldehyde was ound in he ca idge om he
s ain 71B a 1.29 mg, which amoun s o 26 mg/L in he 50-mL olume,
while epo ed le els o his compound in wine a e well below 0.1 mg/L
(Culle ´
e, Cacho, & Fe ei a, 2007). This sugges s ha isobu y aldehyde
is a majo e men a ion ola ile which is nea ly comple ely (>99%) los
by e apo a ion. Simila conside a ions applied o isop opyl and isoamyl
ace a es and e hyl p opanoa e indica e ha mo e han 70% o hese
a omas a e los by e apo a ion. Le els pu ged ou o highe es e s, such
as e hyl bu y a e, isobu y a e, hexanoa e, oc anoa e and decanoa e, and
o highe aldehydes, such as 2 and 3-me hylbu anal, we e mo e modes .
Ye , hey ep esen signi ican ac ions o he o al o med. In he cases
o e hyl hexanoa e and decanoa e, o ins ance, he ac ions los a e 68
and 35% o he o al amoun s o med, espec i ely.
In any case, esul s e eal ha he CO
2
eleased du ing e men a ion
ca ied ou la ge amoun s o isobu y aldehyde, isop opyl ace a e, e hyl
p opanoa e and isoamyl ace a e, which in ac a e mos ly los in his
pe iod; and also signi ican amoun s in absolu e bu no in ela i e
e ms, o isobu anol and isoamyl alcohol. Some o he es e s, such as
e hyl bu y a e, hexanoa e, oc anoa e and decanoa e we e p oduced a
much smalle le els bu ye , we e signi ican ly los by e apo a ion.
3.3. Majo e men a i e a oma compounds
Twen y-six majo e men a ion ola iles we e de ec ed a concen-
a ions supe io o de ec ion limi s in young wines (Table S2). In his
case, i een ou o he wen y-six quan i ied ola iles we e signi ican ly
ela ed o he yeas s ain. The mos di e en p o iles o ola iles we e
ob ained in wines e men ed wi h IONYS, which p oduced maxima
le els o mos e hyl es e s and ace a es, and also o iso ale ic acid,
ace oin and γ-bu y olac one, and minima le els o ace ic and decanoic
acids. O he s ains showing speci ic p o iles o ola iles we e 71B, BDX,
D80, and PERSY. 71B p oduced maxima le els o me hionol, bu anol,
Table 2
Amoun s (in
μ
g) o ola ile compounds pu ged ou du ing e men a ion and apped in LiCh olu -EN ca idges placed be o e he Mulle al es o he e men e s
con aining 50 mL o syn he ic mus . The i s column gi es he signi icance o he ac o yeas on he le els o ola iles ound in he e men a ions ealized wi h PAF
addi ion (p alues in bold a e in e io o 0.05).
p alue MUST CLOS IONYS 71B BDX D254 D80 HPS OKAY PERSY RHONE
Ace a es
p opyl ace a e 6.24E-
10
0.10 ±
0.02
0.6 ±0.2 4.1 ±0.3 0.9 ±0.2 0.49 ±
0.06
0.6 ±0.2 0.5 ±0.2 0.9 ±
0.2
2.1 ±0.4 3.7 ±0.8 0.9 ±0.2
isop opyl ace a e 5.80E-
05
1 ±7 135 ±11 320 ±43 158 ±34 135 ±33 130 ±17 129 ±54 162 ±
43
199 ±18 250 ±33 171 ±23
isobu yl ace a e 4.47E-
02
n.d. 0.7 ±0.1 6 ±2 2.0 ±0.7 1.8 ±0.5 1.2 ±0.4 1 ±1 2.2 ±
0.6
1.4 ±0.4 3.1 ±0.1 1.1 ±0.5
isoamyl ace a e 4.70E-
04
15 ±4 6 ±3 104 ±18 23 ±17 – 17 ±11 8 ±15 18.2 ±
8.1
29 ±5 – 14 ±31
Acids
3-me hylbu y ic
acid
1.14E-
01
n.d. 3 ±1 4 ±3 4 ±2 2.4 ±0.2 4 ±1 1.8 ±0.4 5.1 ±
2.0
2.7 ±3.0 3 ±2 4 ±1
2-me hylbu y ic
acid
7.92E-
02
0.1 ±0.1 0.4 ±0.2 0.8 ±0.7 0.9 ±0.6 0.59 ±
0.09
0.9 ±0.4 0.41 ±
0.09
1.1 ±
0.5
1.4 ±0.8 0.6 ±0.6 1. 0 ±0.3
Alcohols
isobu anol 4.69E-
02
74 ±20 130 ±24 129 ±8 128 ±4 – 176 ±30 142 ±
112
184 ±
31
127 ±54 – 120 ±16
isoamyl alcohol 1.57E-
01
387 ±13 420 ±
107
488 ±58 484 ±
226
– 497 ±
163
325 ±
318
507 ±
125
664 ±
116
– 364 ±
110
Ca bonyls
ace aldehyde 3.07E-
03
5 ±2 3 ±2 3.6 ±0.6 14 ±10 – 3.3 ±0.8 4 ±1 4 ±1 4 ±2 – 5 ±2
isobu y aldehyde 6.39E-
02
n.d. 904 ±
482
684 ±
124
1291 ±
528
957 ±
464
548 ±
268
392 ±
210
692 ±
367
493 ±
267
644 ±
243
534 ±
113
2-me hylbu anal 5.85E-
02
n.d. 1.4 ±0.2 2.2 ±0.7 3 ±1 1.1 ±0.5 1 ±1 0.7 ±0.1 1.2 ±
0.8
2.4 ±0.8 1.7 ±0.1 0.9 ±0.9
3-me hylbu anal 2.05E-
01
n.d. 3.5 ±0.9 7 ±1 12 ±4 4.2 ±2.3 3 ±4 3. 0 ±
0.3
4 ±3 10 ±4 6.5 ±0.3 4 ±4
Es e s
e hyl p opanoa e 5.28E-
04
0.1 ±0.2 6.5 ±0.6 38 ±18 5 ±1 4.9 ±0.3 6 ±7 5 ±2 7 ±2 7.5 ±0.5 10 ±3 6 ±2
e hyl bu y a e 2.95E-
02
0.2 ±0.1 0.10 ±
0.03
0.86 ±
0.05
0.19 ±
0.01
– 0.23 ±
0.04
0.1 ±0.1 0.3 ±
0.1
0.20 ±
0.03
– 0.2 ±0.1
e hyl isobu y a e 3.07E-
03
n.d. 0.3 ±0.2 0.30 ±
0.05
0.32 ±
0.01
0.34 ±
0.15
0.5 ±0.2 0.21 ±
0.09
0.5 ±
0.1
0.32 ±
0.06
0.38 ±
0.02
0.4 ±0.1
e hyl hexanoa e 3.09E-
01
2.7 ±0.8 4 ±2 7.0 ±0.6 6 ±5 – 5.9 ±2.3 3 ±1 4 ±2 6.7 ±0.9 – 3 ±5
e hyl oc anoa e 4.89E-
01
10 ±5 7 ±4 11 ±2 11 ±8 – 11 ±4 5 ±1 7 ±3 14 ±4 – 6 ±9
e hyl decanoa e 1.23E-
04
0.32 ±
0.05
2.2 ±0.7 8 ±2 4 ±2 – 4 ±1 2.9 ±0.9 3.6 ±
0.9
6.3 ±0.1 – 5 ±1
n.d. indica es ha he compound was no de ec ed o below de ec ion limi s.
– indica es ha da a is no a ailable.
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
5
hexanol and minima le els o mos e hyl es e s. BDX p oduced maxima
le els o isobu anol and isoamyl alcohol. D80 p oduced maxima le els o
ace ic, isobu y ic and decanoic acids. Finally, PERSY p oduced maxima
le els o e hyl oc anoa e, oc anoic acids, e hyl lac a e and minima le els
o isoamyl alcohol and isobu anol.
In gene al, di e ences in oduced by he s ains we e o li le o
mode a e magni ude. Le els o isoamyl alcohol anged om 160 o 310
mg/L, a ac o 2; hose o isobu anol om 20 o 60 mg/L, a ac o 3.
These di e ences a e, howe e , la ge enough o ha e senso y signi i-
cance (De-la-Fuen e-Blanco, S´
aenz-Na ajas, & Fe ei a, 2017). Di e -
ences in he le els o ace ic acid we e much highe and amoun ed o a
ac o 20, om jus 30 mg/L (IONYS) o 600 mg/L (RHONE). Howe e ,
lea ing aside IONYS, di e ences become mo e modes , anging om
340 o 600 mg/L, less han a ac o 2. Simila anges o a ia ion we e
obse ed o hexanoic, oc anoic, decanoic and iso ale ic acids, while
le els o isobu y ic acid anged a ac o close o 4. Le els o es e s we e,
in gene al, e y low as a possible consequence o hei s ong e apo a-
ion, and in some cases, hey we e no e en de ec ed. Lea ing aside
IONYS, hei anges o a ia ion we e no la ge.
3.4. T ace a oma compounds. Va ie al o e men a i e o igin?
The comple e da a se s wi h he concen a ions o up o hi y- ou
ace a oma componen s in ecen ly e men ed and in aged wines, in
he di e en con ols in oduced in he s udy, and he esul s o he
di e en ANOVA s udies ca ied ou on he da a a e compiled in he
Supplemen a y ma e ial (Tables S2–S6).
Rega ding he a ie al o e men a i e o igin o he a oma com-
pounds, he s udy o he con ols including o no he PAF ma e ial
ex ac ed om he g apes and hose o he s including o no e men a ion
(all compiled in Table S3), e eals ha some a oma compounds canno
be unequi ocally classi ied in o e men a i e o a ie al. Ra he , he e
a e se e al in e media e ca ego ies, as he answe o he ollowing h ee
simple ques ions asked o each a oma compound, e eals;
1. Is he a oma compound p esen in e men ed con ols no con aining
g ape PAF?
2. Is i p esen in un e men ed con ols con aining g ape PAF?
3. Is i a signi ican ly la ge amoun s in e men ed samples con aining
g ape PAF han in he co esponding con ols no con aining g ape
PAF?
A posi i e answe o he i s ques ion implies a unequi ocal
e men a i e o igin; he compound can be o med by yeas s om he
basic lis o nu ien s supplied. On he con a y, a nega i e answe in-
dica es ha he o ma ion o he compound equi es he p esence o
g ape componen s. The answe s o he wo ollowing ques ions will
indica e whe he he compound is p esen in he g ape PAF as speci ic
p ecu so (posi i e answe o 2) and whe he yeas is equi ed o i s
o ma ion (posi i e answe o 3). A ending o he answe s, i e di e en
o igin- ela ed ca ego ies eme ged, as is schema ized in Fig. 1.
1) Pu e e men a i e compounds (answe s YNN) a e hose which we e
p esen in e men ed samples no con aining PAF and whose le els
we e no in luenced by he p esence o PAF. Compounds in his
ca ego y we e isobu yl ace a e, e hyl iso ale a e, e hyl 2-me hylbu-
y a e and γ-decalac one.
2) PAF-modula ed e men a i e compounds (answe s YNY) a e hose
a oma compounds o med by yeas , bu whose le els a e signi i-
can ly in luenced by he p esence o PAF. Compounds in his ca e-
go y we e β-phenyle hyl ace a e, e hyl iso ale a e, e hyl leuca e,
γ-oc alac one, β-ci onellol, ge aniol, ne ol.
3) Fe men a i e and a ie al a oma compounds (answe s YYY and
YYN), a e hose a oma compounds which can be o med by yeas
om basic nu ien s and which can be also ound in un e men ed
con ols con aining PAF. Linalool and linalool oxide belong o his
ca ego y.
4) Yeas -induced a ie al a oma compounds (answe s NNY), which a e
hose a oma compounds ound exclusi ely in e men ed PAF. E hyl
dihyd ocinnama e and β-ionone belong o his ca ego y.
5) Va ie al a oma compounds (answe s NYY), which a e hose com-
pounds ound exclusi ely in samples con aining g ape PAF. Mos
compounds belong o his ca ego y (massoia lac one, β-dam-
ascenone, TDN, i ispi ane, Riesling ace al, anillin, ace o anillone,
sy ingaldehyde, sy ingol, guaiacol, 4-e hyguaiacol, 4-e hylphenol, 4-
inylguaiacol, 4- inylphenol, eugenol, me hoxyeugenol, ans-iso-
eugenol, p-p opylguaiacol).
I should be no ed ha , di ec ly o indi ec ly, le els o all compounds
we e in luenced by he exis ence o e men a ion, which sugges s, as i
will be u he seen in he nex sec ions, ha yeas is going o play a
ele an ole on nea ly he comple e wine a oma p o ile.
3.5. Yeas s ain and aging: global o e iew
Bo h ac o s, yeas s ain and aging ha e a s ong in luence on he
ace a oma composi ion o wines, al hough aging is he dominan ac-
o . Such dominance is mos e iden in he PCA plane gi en in Fig. 2,
which shows he p ojec ion o he 60 samples (10 yeas s ×2 imes o
analysis ×3 eplica es) in he wo i s dimensions. The i s componen
(55.1% o he o iginal a iance) sepa a es samples a ending o age,
wi h young samples on he le , and aged samples on he igh . The
a iable loadings (no shown) e eals ha mos compounds, including
no isop enoids, es e s, ola ile phenols, anillin de i a i es, inc ease
du ing aging and ha only e penes (excep o linalool oxide), massoia
lac one and phenyle hyl ace a e dec ease du ing aging.
The plo e eals wo o he impo an cha ac e is ics. Fi s , ha
samples e men ed wi h IONYS a e e y well sepa a ed om he o he s
be o e and a e aging. Young samples because o i s highes le els in
β-phenyle hyl ace a e, linalool and ge aniol, and aged samples because
o i s highes le els o isobu yl ace a e, γ-oc a and decalac one. The
second ema kable cha ac e is ic, is ha lea ing aside IONYS, he yeas
s ain is an ac i e g ouping ac o in ace a oma compounds only a e
aging.
3.6. E ec s o yeas s ain on wine a oma
In o de o analyse he in luence o yeas aking in o conside a ion
he dominance o he aging ime, wo di e en hea maps we e gene a ed
wi h a oma compounds signi ican ly a ec ed by yeas s. The i s is gi en
in Fig. 3A and includes da a om majo and ace a oma compounds
Fig. 1. The i e o igin- ela ed ca ego ies in which a oma compounds o he
expe imen should be classi ied.
M. Dena e al.

Food Chemis y: X 9 (2021) 100116
6
measu ed in young wines. The second can be seen in Fig. 3B and includes
only ace a oma compounds in aged wines (Fig. 3B). The esul s o he
hie a chical clus e ings a e also displayed on he le pa o each plo .
Fig. 3A con i ms he singula i y o IONYS in young wines and he
appa en ly low di e si y exis en be ween he o he s ains. A e aging,
howe e , a much clea s uc u e eme ges as can be seen in Fig. 3B,
whe e yeas s ains can be classi ied in o h ee di e en clus e s. Clus e
1 is in eg a ed by a qui e homogeneous g oup o med by D254, HPS,
RHONE and CLOS and wi h mo e dissimila i y, by D80. The second
clus e is o med by PERSY, OKAY, 71B and wi h mo e dissimila i y,
BDX. And inally, IONYS is he single componen o he mos di e en
clus e 3. I can also be seen ha yeas s in clus e 1 p oduced highe
le els o pu e e men a i e compounds such as isobu yl ace a e, e hyl
isobu y a e, 2-me hylbu y a e and iso ale a e and eleased highe le els
o no isop enoids (TDN, i ispi ane and Riesling ace al). Clus e 2
eleased in gene al smalle le els o ola iles, excep o some ola ile
phenols, such as anillin and guaiacol.
3.7. Modula ion o a ie al a oma
The e ec o yeas s ain on hose genuine a ie al a oma compounds
na u ally p esen in un e men ed con ols con aining g ape PAF can be
assessed wi h he help o he plo s gi en in Fig. 4. The plo s compa e
le els o a oma compounds ound in e men ed aged samples wi h hose
ob ained in he un e men ed aged con ols. These ep esen a ions
acili a e he iden i ica ion o he gene al ole o e men a ion on he a e
o hese a ie al a oma compounds and also o he speci ic ole played
by he s ain o yeas . Compounds can be classi ied in o h ee di e en
ca ego ies depending on he e ec o e men a ion.
1s ca ego y.- Posi i e e ec o e men a ion. This ca ego y includes
i e a oma compounds whose le els in e men ed samples we e much
abo e hose ound in he un e men ed con ols, sugges ing ha some o
he speci ic p ecu so s o hese a oma compounds could be o med by
he ac ion o yeas s. Compounds in his ca ego y a e ace o anillone,
β-ionone ( his one is no shown in Fig. 4, since i was only ound in young
samples), e hyl dihyd ocinnama e, p-p opylguaiacol, 4-e hylphenol,
ans-isoeugenol ( hese ou a e no ep esen ed in Fig. 4 since hey
we e no de ec ed in he un e men ed con ol) and eugenol. Only in he
case o eugenol he e ec o yeas was signi ican . Le els o ace o a-
nillone in aged e men ed samples a e 10 imes highe han hose ound
in he un e men ed con ol, howe e , no di e ence was obse ed be-
ween he yeas s.
2nd ca ego y.- No e ec o e men a ion. Compounds in his ca ego y
ha e in common ha , in a e age, le els o e men ed samples a e no
dissimila o hose o he un e men ed con ols. The e a e howe e
s ong di e ences a ending o he di e en ial e ec in oduced by
yeas . In he cases o Riesling ace al in Fig. 4A, and o me hoxyeugenol
and β-damascenone in Fig. 4B, he e is no e ec o yeas , which sugges s,
Fig. 2. PCA ca ied ou on concen a ions o ace a oma compounds in he PAF-con aining samples e men ed by 10 S. ce e isiae s ains, analysed a e e men a ion
and a e accele a ed aging.
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
7
ha a leas o Temp anillo, he le els o hese a ie al a oma com-
pounds canno be modula ed by yeas . The case o β-damascenone,
which is an a oma enhance (Pineau, Ba be, Van Leeuwen, & Dubou -
dieu, 2007) and modules he ipeness-cha ac e o ui y pe cep ion
(San-Juan, Fe ei a, Cacho, & Escude o, 2011), dese es special
men ion. Le els o his odo an in ecen ly e men ed samples we e
abo e hose measu ed in he co esponding un e men ed con ols
(Supplemen a y ma e ial S2 and S5) and we e signi ican ly in luenced
by he s ain o yeas . Samples e men ed wi h IONYS showed le els up
o 4 imes highe han hose ound in hose made wi h OKAY. This
sugges s ha yeas s canno change he long- e m le el o his a oma
compound bu can accele a e i s o ma ion. The second subca ego y
includes anillin (4C), me hoxyeugenol (4B), and 4- inylphenol (4D),
o which he s ain o yeas exe s a mode a e and signi ican in luence,
so ha di e ences o a ound a 50% be ween he minimum and he
maximum a e obse ed. 4-Vinylguaiacol can be also classi ied wi hin
his subca ego y bu wi h wo s ains, IONYS and OKAY, showing a clea
ou lie o e p oduc i e cha ac e .
3 d ca ego y.- Nega i e e ec o e men a ion. This ca ego y in-
cludes a oma compounds whose le els in e men ed samples we e below
hose ound in he un e men ed con ols. The e a e s ong di e ences
be ween compounds a ending o he e ec played by he s ain o yeas .
Massoia lac one (Fig. 4B) is a pa icula case whose le els d op o nea ly
ze o in all e men ed samples wi h no di e ence be ween s ains. Mas-
soia lac one is an impo an ma ke o o e - ipeness and con ibu o o
p une a oma, whose le els a e known o dec ease du ing e men a ion
(Pons, Allamy, La igne, Dubou dieu, & Da ie , 2017). Ou esul s
e eal ha such dec ease in ensi ies du ing aging, which sugges s ha
e men a ion educes also he p ecu so s. I would be o in e es o see
whe he such educ ion is equally e ec i e in g apes con aining highe
le els o p ecu so s o his molecule. Compounds in he ca ego y o
which he s ain o yeas in oduced signi ican di e ences we e, om
less o mo e in ense he e ec o s ain: sy ingol, i ispi ane, sy ingal-
dehyde, guaiacol and TDN. I is appa en ha he speci ic p ecu so s o
hese compounds a e me abolized di e en ly by he di e en s ains.
This can ha e s ong echnological ele ance since guaiacol and TDN
can ake pa in ele an odou aul s, and sugges s ha selec ed s ains
a e a po en ially e ec i e emedial ool. Guaiacol is an a oma compound
con ibu ing o he cha ac e is ic oas y-woody no es o Temp anillo,
bu i can be a se ious o -odou de eloped wi h ime in wines made wi h
g apes exposed o smoke (Ris ic, Van De Huls , Capone, & Wilkinson,
2017). Resul s in Fig. 4 e eal ha yeas s wi hin he clus e 1 seem o
me abolize he p ecu so a highe le els. In he case o RHONE, le els o
guaiacol we e educed by almos a ac o 3 compa ing wi h he un e -
men ed con ol. Powe ul educ ions linked o speci ic yeas s ains can
be also obse ed o TDN, known esponsible o ke osene no es de el-
oped in aged Riesling wines. Wi h a 2
μ
g/L de ec ion h eshold (Sacks
e al., 2012), his compound will su ely also con ibu e o unpleasan
no es in aged ed wines. The igu e also e eals ha yeas s in clus e 2,
no ably 71B can TDN educe le els by a ac o 3 wi h espec o he
con ol o D80. A simila yeas -induced and i ispi ane-independen
dec ease o TDN le els has been ecen ly obse ed o non-Saccha o-
myces yeas (Oli ei a & Fe ei a, 2019) bu o he bes o ou knowledge,
i has no been obse ed o Saccha omyces s ains. This abili y can ha e
a no able senso y impo ance, since le els o TDN a e expec ed o in-
c ease due o clima e change (Win e hal e & G¨
ok, 2013).
Fig. 3. Hea maps ob ained om no malized da a o he main ola iles signi ican ly modula ed by yeas s in young (A) and aged (A) wines. Compounds a e de ailed in
he sub igu e C.
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
8
Many a ie al a oma compounds in Fig. 4 de i e om e ulic acid
and i s glycosides ( anillin, ace o anillone, isoeugenol, eugenol and 4-
inylguaiacol). The highe le els o hese a oma molecules measu ed
in aged e men ed samples sugges ha yeas ans o ms e ulic acid
glycosides in o he co esponding a oma glycosides and ha hose
ans o ma ions a e s ain speci ic. Aged wines made wi h BDX ha e
maxima con en s o anillin and ace o anillone, hose made wi h IONYS
ha e maxima con en s o eugenol while hose made wi h OKAY ha e
minima le els o anillin and maxima le els o 4- inylguaiacol. Some o
hose speci ici ies seem o be sha ed by s ains in he same clus e . Yeas s
in clus e 2 show highe le els o anillin, ace o anillone and 4- inyl-
guaiacol han hose in clus e 1. 4-Vinylguaiacol and 4- inylphenol
dese e a speci ic commen , since hei le els a e ex emely dependen
o he yeas s ain in unaged samples, wi h ac o s a ound 10 be ween
he minimum and maximum concen a ions. Because o hei eac i i y,
di e ences be ween maxima and minima sh ink o ac o s 3 (case o 4-
inylguaiacol) and 1.7 (case o 4- inylphenol). In bo h cases, maxima
le els we e obse ed o IONYS, and minima le els o BDX and HPS.
3.8. Modula ion o o he ele an a oma molecules
Linalool and ge aniol a e he wo mos impo an e penols o wine.
In he p esen case, hese wo molecules we e ha dly de ec ed in he
un e men ed con ols, sugges ing ha he g ape ma e ial did no ha e
much p ecu so s. The wo compounds we e ound in he con ols no
con aining g ape ex ac , so ha yeas s we e able o o m weak, o
mode a e in he case o IONYS, amoun s o hese molecules. Fe men ed
unaged samples con ained bo h molecules a he expec ed concen a ion
anges o Temp anillo (<6
μ
g/L) excep samples e men ed by IONYS,
whose le els we e abo e 20
μ
g/L, which sugges s ha his qui e unique
yeas s ain will p oduce young wines wi h ma kedly di e en cha ac-
e s. Du ing aging, le els o hese compounds and he o he e penes
dec eased (<1
μ
g/L), while hose o linalool oxide, i s oxida ion p oduc ,
inc eased.
E hyl leuca e o e hyl 2-hyd oxy-4-me hyl ale a e, is a ema kable
a oma compound iden i ied in aged wines (Campo, Cacho, & Fe ei a,
2006) and sugges ed o be key in he speci ic blackbe y a oma o
Bo deaux ed wines (Falcao, Ly a, Da ie , & Ba be, 2012). Resul s om
his pape ha e shown ha maxima le els a e ound in aged PAF-
con aining e men ed samples and ha he yeas s ain exe s a signi i-
can in luence, wi h le els ound in IONYS, BDX, D80, PERSY and
RHONE wice hose ound in 71B o OKAY (Table S5).
E hyl es e s o b anched acids a e he mos impo an ui y es e s in
aged ed wines, con ibu ing conce edly o ui y a oma (De-la-Fuen e-
Blanco, S´
aenz-Na ajas, Valen in, & Fe ei a, 2020) and a e slowly
o med by es e i ica ion o he co esponding acids syn hesized du ing
e men a ion h ough Eh lich pa hway (Swiege s, Ba owsky, Henschke,
& P e o ius, 2005). The in luence o yeas becomes mos ob ious a e
aging. Minima le els a e ound in 71B, PERSY (clus e 2), and maxima in
D80 and D254 (clus e 1), wi h di e ences as la ge as ac o s be ween 4
and 6.5.
Isoamyl and phenyle hyl ace a es a e ele an in he a oma o young
wines, since hei le els quickly ade by hyd olysis o he es e s. The
in luence o yeas s in hei le els is o e whelming. In he case o
β-phenyle hyl ace a e, le els in young wines (Table 5, Supplemen a y
ma e ial) ange om 0.1 mg/L (clus e 1) o 1.5 mg/L in clus e 3, wi h
le els in samples om clus e 2 be ween 0,18 and 0.29 mg/L. Simila
esul s we e al eady obse ed o he isoamyl ace a e and isop opyl
ace a es e apo a ed du ing e men a ion (Table 2).
4. Conclusions
Vola iles los by e apo a ion du ing e men a ion a e mos ly
e men a i e compounds and no g ape- ela ed a oma compounds.
Quan i a i ely, apou s a e majo ly composed o 2-me hylp opanal,
isop opyl ace a e, isoamyl ace a e, e hyl p opanoa e, isobu anol and
isoamyl alcohol. While he ac ion o alcohols los is e y low, ha o
he aldehydes and es e s can be well abo e 90% o he o al ola ile
Fig. 4. Concen a ions o a ie al a oma compounds in PAF-con aining aged e men ed samples acco ding o hei le els in he un e men ed con ol.
M. Dena e al.
Food Chemis y: X 9 (2021) 100116
9
p oduced.
The s ong impac exe ed by he s ain o yeas on wine a oma
composi ion becomes in many cases only e iden a e aging, since
le els o e hyl es e s o b anched acids, o mos g ape- ela ed a oma
compounds and o many mino yeas s-de i ed a oma compounds mos ly
inc ease du ing aging. The 10 s ains can be classi ied in o h ee clus e s
showing ma ked di e ences in e men a i e and a ie al a oma p o iles.
The bounda ies be ween e men a i e and a ie al a oma com-
pounds a e in many cases blu ed. Fi s , he s udy has con i med a
e men a i e o igin o linalool and ge aniol, ound a high le els in
samples e men ed by one o he s ains. Second, he p esence o poly-
phenolic and a oma ic ac ions om g ape exe s a s ong in luence on
yeas me abolism and, hi d, he s ains o yeas no only hyd olyze
glycosidic p ecu so s, bu me abolize qui e di e en ly he p ecu so s o
ele an a oma compounds, such as phenolic acids and no isop enoids.
These cha ac e is ics ha e in e es ing p ac ical consequences on he
po en ial o yeas s o con ol he wine a oma p o ile and, mos
ema kably, some wine aging a ibu es. Resul s ha e shown ha he
a es o accumula ion o β-damascenone a e s ain- ela ed, and ha
some s ains may be speci ically used o mi iga e ele an aging- ela ed
o -odou s, such as hose ela ed o guaiacol, massoia lac one o TDN.
CRediT au ho ship con ibu ion s a emen
Ma ie Dena : In es iga ion, Fo mal analysis, Da a cu a ion, W i ing
- o iginal d a , Visualiza ion. Dolo es P´
e ez: In es iga ion, W i ing -
e iew & edi ing. Jos´
e Ma ía He as: Concep ualiza ion, Resou ces,
W i ing - e iew & edi ing. Ampa o Que ol: Supe ision, Me hodology,
W i ing - e iew & edi ing. Vicen e Fe ei a: Supe ision, Valida ion,
Me hodology, W i ing - e iew & edi ing, Visualiza ion.
Decla a ion o Compe ing In e es
The au ho s decla e ha hey ha e no known compe ing inancial
in e es s o pe sonal ela ionships ha could ha e appea ed o in luence
he wo k epo ed in his pape .
Acknowledgemen s
The p ojec leading o his applica ion has ecei ed unding om he
Eu opean Union’s Ho izon 2020 esea ch and inno a ion p og amme
unde he Ma ie Skłodowska-Cu ie g an ag eemen No 764364. Ana
Escude o is acknowledged o he supply o Temp anillo g apes and Sa a
Fe e o-Del-Teso o he p epa a ion o Temp anillo mis ela. The au-
ho s decla e ha no con lic o in e es exis s.
Appendix A. Supplemen a y da a
Supplemen a y da a o his a icle can be ound online a h ps://doi.
o g/10.1016/j. ochx.2021.100116.
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