2021
125
Inés Pe ei a Biscaia de Oli ei a
Con ibu ion o wine
mic oo ganisms o he a oma
composi ion o wine and i s
senso y impac
Di ec o /es
Fe ei a González, Vicen e
Fische , Ul ich
© Uni e sidad de Za agoza
Se icio de Publicaciones
ISSN 2254-7606
Inés Pe ei a Biscaia de Oli ei a
CONTRIBUTION OF WINE MICROORGANISMS TO
THE AROMA COMPOSITION OF WINE AND ITS
SENSORY IMPACT
Di ec o /es
Fe ei a González, Vicen e
Fische , Ul ich
Tesis Doc o al
Au o
2019
Reposi o io de la Uni e sidad de Za agoza – Zaguan h p://zaguan.uniza .es
UNIVERSIDAD DE ZARAGOZA
Escuela de Doc o ado
P og ama de Doc o ado en Ciencia Analí ica en Química
Con ibu ion o wine mic oo ganisms o he
a oma composi ion o wine and i s senso y
impac
PhD Thesis by
Inês Pe ei a Biscaia de Oli ei a
Janua y 2019
Supe iso s:
D . Vicen e Fe ei a González
D . Ul ich Fische

2
Con ibu ion o wine mic oo ganisms o he
a oma composi ion o wine and i s senso y
impac
by
Inês Pe ei a Biscaia de Oli ei a
Disse a ion p esen ed o he deg ee o
Doc o o Philosophy in Analy ical Chemis y
Janua y 2019
Supe iso s:
D . Vicen e Fe ei a González
D . Ul ich Fische
D. VICENTE FERREIRA GONZÁLEZ, Ca ed á ico del Depa amen o de
Química Analí ica y D. ULRICH FISCHER, di ec o en el Ins i u o DLR-
Rheinp alz, Alemania
CERTIFICAN:
Que la p esen e memo ia, i ulada “Con ibu ion o wine mic oo ganisms
o he a oma composi ion o wine and i s senso y impac ” co espondien e al
plan de in es igación ap obado po la Comisión de Doc o ado del Depa amen o
de Química Analí ica y p esen ada pa a op a al g ado de doc o a en Ciencia
Analí ica en Química, ha sido ealizada bajo nues a di ección po Dª. Inês
Pe ei a Biscaia de Oli ei a, au o izando su p esen ación pa a p osegui los
ámi es opo unos y p ocede a su cali icación po el ibunal co espondien e.
Za agoza, 17 ene o de 2019.
Fdo. D . Vicen e Fe ei a González Fdo. D . Ul ich Fische

Index
Index
Index
PRESENTATION ................................................................................................................ 3
INTRODUCTION ................................................................................................................ 7
1. WINE FLAVOUR AND ITS PERCEPTION .............................................................................. 9
1.1 Wine a oma genesis............................................................................................... 10
1.2 G ape a ie y- ela ed non-speci ic p ecu so s ....................................................... 12
1.3 Glycosidic p ecu so s ............................................................................................ 13
1.4 Cys einyla ed and glu a hionyla ed p ecu so s ....................................................... 14
1.5 E ec s o ex e nal addi i es on wine a oma ........................................................... 15
2. WINE AROMA FORMATION FROM SPECIFIC AND NON-SPECIFIC PRECURSORS DURING
ALCOHOLIC FERMENTATION .............................................................................................. 16
2.1 Spon aneous e sus inocula ion e men a ions ....................................................... 17
2.2 Yeas me abolism du ing alcoholic e men a ion .................................................... 18
2.3 Release o a oma compounds om glycosidic p ecu so s ....................................... 22
2.3.1 Release and o ma ion o mono e penes ................................................................. 22
2.3.2 Release and o ma ion o no isop enoids................................................................ 24
2.3.3 Release and o ma ion o ola ile phenols .............................................................. 26
3. SELECTION OF NON-SACCHAROMYCES YEAST STRAINS .................................................. 27
3.1 To ulaspo a delb ueckii ........................................................................................ 28
3.2 Pichia kluy e i ...................................................................................................... 28
3.3 Lachancea he mo ole ans..................................................................................... 28
4. BIBLIOGRAPHY ............................................................................................................. 30
OBJECTIVES ................................................................................................................... 39
SECTION I ........................................................................................................................ 41
STRECKER ALDEHYDES ARE NORMAL BY-PRODUCTS OF ALCOHOLIC
FERMENTATION LINKED TO YEAST SULPHITE METABOLISM
1. INTRODUCTION ............................................................................................................. 43
2. METHODOLOGY ............................................................................................................ 46
2.1 Reagen s and s anda ds ......................................................................................... 46
2.2 Cul u e condi ions: ................................................................................................ 46
Index
2.2.1 Syn he ic mus composi ion ...................................................................................46
2.2.2 Yeas cul u e and e men a ion se -up.....................................................................47
2.3 Analy ical me hods ................................................................................................ 48
2.3.1 Classical oenological cha ac e iza ion ....................................................................48
2.3.2 Quan i ica ion o o al aldehydes ............................................................................49
2.3.3 Analysis o majo ola ile compounds ....................................................................50
2.4 Da a ea men ...................................................................................................... 50
3. RESULTS AND DISCUSSION ............................................................................................ 53
3.1 S ecke aldehydes a e no mal e men a i e compounds ........................................ 55
3.2 Role o SO2 ........................................................................................................... 60
3.3 Role o Zn on SA o ma ion ................................................................................... 63
3.4 Fa y acids and hei e hyl es e s ........................................................................... 63
4. CONCLUSIONS .............................................................................................................. 66
5. BIBLIOGRAPHY ............................................................................................................ 68
SECTION II ....................................................................................................................... 73
ROLES OF YEAST ON THE FORMATION AND EVOLUTION OF THE AROMA OF
RIESLING AND GARNACHA WINES
INTRODUCTION AND METHODOLOGY .................................................................... 75
1. SECTION II – INTRODUCTION ........................................................................................ 77
2. GOALS ......................................................................................................................... 80
3. SECTION II – METHODOLOGY ....................................................................................... 81
3.1 Reagen s and s anda ds ........................................................................................ 81
3.2 Glycosidic p ecu so s ex ac ion ........................................................................... 81
3.2.1 G ape p ocessing ...................................................................................................81
3.2.2 Glycosidic ex ac ion using a SPE based me hod ....................................................82
3.3 Syn he ic mus e men a ion................................................................................... 83
3.3.1 Syn he ic mus composi ion ...................................................................................83
3.3.2 Yeas cul u e .........................................................................................................84
3.3.3 Fe men a ion se -up and accele a ed aging .............................................................84
3.3.4 Un e men ed con ols ............................................................................................85
3.4 Analy ical me hods ................................................................................................ 86
3.4.1 Analysis o mino and ace ola ile compounds .....................................................86
3.4.2 Ch oma og aphic me hod.......................................................................................86
Index
3.5 Da a ea men ...................................................................................................... 89
4. BIBLIOGRAPHY ............................................................................................................. 90
CHAPTER 1 ....................................................................................................................... 95
EFFECTS OF SEQUENTIAL INOCULATION WITH DIFFERENT NON-
SACCHAROMYCES ON THE FORMATION AND FURTHER EVOLUTION OF
RIESLING WINE AROMA
1. RESULTS AND DISCUSSION ............................................................................................ 97
1.1 Classic oenological pa ame e s o inal syn he ic wines ......................................... 98
1.2 O e iew o he a oma composi ion o inal wines .................................................. 99
1.3 Fe men a i e compounds in Riesling.................................................................... 102
1.4 Va ie al compounds in Riesling ........................................................................... 110
2. CONCLUSIONS ............................................................................................................ 121
3. BIBLIOGRAPHY ........................................................................................................... 122
CHAPTER 2 ..................................................................................................................... 127
EFFECTS OF SEQUENTIAL INOCULATION WITH DIFFERENT NON-
SACCHAROMYCES ON THE FORMATION AND FURTHER EVOLUTION OF
GARNACHA WINE AROMA
1. RESULTS AND DISCUSSION .......................................................................................... 129
1.1 Classical oenological pa ame e s o inal syn he ic wines .................................... 130
1.2 O e iew o he a oma composi ion o inal wines ................................................ 131
1.3 Fe men a i e compounds in Ga nacha wines ....................................................... 135
1.4 Conside a ions abou po en ial senso y e ec s o di e en yeas s ains ............... 142
1.5 Va ie al compounds in Ga nacha wines ............................................................... 144
2. CONCLUSIONS ............................................................................................................ 153
3. BIBLIOGRAPHY ........................................................................................................... 156
CHAPTER 3 ..................................................................................................................... 159
OBSERVATIONS, QUESTIONS AND CONCLUSIONS DERIVED FROM THE
COMPARISON BETWEEN VARIETIES
1. OBJECTIVE ................................................................................................................. 161
2. RESULTS AND DISCUSSION .......................................................................................... 161

Index
2.1 No isop enoids o ma ion and e olu ion in Riesling and Ga nacha...................... 164
2.2 Mono e penes in Riesling and Ga nacha ............................................................. 172
2.3 Vola ile phenols in Riesling and Ga nacha .......................................................... 175
2.4 Medium chain a y acids e hyl es e s in Riesling and Ga nacha .......................... 177
2.5 Fusel alcohols in Riesling and Ga nacha ............................................................ 178
3. CONCLUSIONS ............................................................................................................ 180
4. BIBLIOGRAPHY .......................................................................................................... 182
SECTION III ................................................................................................................... 185
AROMA OF GERMAN RIESLING WINES AND INFLUENCE OF THE TERROIR 185
INTRODUCTION AND METHODOLOGY
1. SECTION III - INTRODUCTION ...................................................................................... 189
2. SECTION III - METHODOLOGY CHAPTER 4 ................................................................... 193
2.1 P ojec collabo a ion .......................................................................................... 193
2.2 Riesling comme cial wines .................................................................................. 193
2.3 Senso y analysis: ................................................................................................ 193
2.4 Ch oma og aphy-ol ac ome y analysis ............................................................... 195
2.5 Vola ile compounds quan i ica ion ...................................................................... 196
2.6 Da a analysis ...................................................................................................... 196
3. SECTION III - METHODOLOGY FROM CHAPTER 5 .......................................................... 197
3.1 P ojec collabo a ion .......................................................................................... 197
3.2 Ha es and wine e men a ion ............................................................................ 197
3.3 Chemical analysis ............................................................................................... 198
3.4 Da a analysis ...................................................................................................... 200
4. BIBLIOGRAPHY .......................................................................................................... 202
CHAPTER 4 .................................................................................................................... 205
SENSORY AND CHEMOSENSORY CHARACTERIZATION OF COMMERCIAL
YOUNG RIESLING WINES FROM DIFFERENT GERMAN APPELLATION
1. RESULTS AND DISCUSSION .......................................................................................... 207
1.1 Senso y Desc ip i e Analysis ............................................................................... 207
1.2 Semiquan i a i e Gas Ch oma og aphy-Ol ac ome y and GC quan i a i e analysis
................................................................................................................................. 211
Index
1.3 In eg a ion o senso y, semiquan i a i e and quan i a i e da a.............................. 215
2. CONCLUSIONS ............................................................................................................ 219
3. BIBLIOGRAPHY ........................................................................................................... 220
CHAPTER 5 ..................................................................................................................... 223
IMPACT OF VINEYARD VERSUS CELLAR MICROBIOTA FROM DIFFERENT
HARVESTS ON THE DISTINCTION OF DIFFERENT RIESLING VINEYARD
1. RESULTS AND DISCUSSION .......................................................................................... 225
1.1 Impo ance o ha es yea e ec and e men a ion loca ion ................................. 225
1.2 Vineya d e sus wine y ....................................................................................... 230
2. CONCLUSIONS ............................................................................................................ 236
3. BIBLIOGRAPHY ........................................................................................................... 238
SUPPLEMENTARY DATA ............................................................................................ 241
1. ODOUR THRESHOLDS .................................................................................................. 243
2. SUPPLEMENTARY DATA FROM SECTION I ..................................................................... 245
3. SUPPLEMENTARY DATA FROM SECTION II: CHAPTER 1 ................................................. 248
4. SUPPLEMENTARY DATA FROM SECTION II: CHAPTER 2 ................................................. 253
5. SUPPLEMENTARY DATA FROM SECTION III: CHAPTER 2 ................................................ 259
P esen a ion

In oduc ion
9
In oduc ion
1. Wine la ou and i s pe cep ion
Fla ou o oods has long been desc ibed as he complex in e ac ion o odou ,
a oma, as e and mou h eel. Odou comp ises he pe cep ion o ola iles di ec ly
by o ho-nasal ia while he a oma en ails he pe cep ion o ola iles which a e
inges ed and de ec ed by e o-nasal ia. Majo ac o s di e en ia ing odou and
a oma a e ola iliza ion empe a u e and he dis inc i e mass ans e condi ions
be ween he p oduc and he p oduc sp ead in buccal mucosa. Tas e is well
known by i s a ibu es swee , sou , sal y, bi e and umami sensed by ongue
ecep o s and mou h eel desc ibes he ac ile sensa ion p oduced by ood (Bae
e al., 2012).
I is no an easy ask o de ine la ou since i depends on some aspec s which
a e speci ic o each indi idual, such as he sensi i i ies o hei ol ac o y and
as e ecep o s and hei deg ee o expe ience. The la ou is ela ed no only
wi h he p esence o as e and odou -ac i e molecules in he p oduc and o hei
concen a ion p o ile, bu also wi h he way in which hose molecules in e ac
wi h he ma ix, o he possible exis ence o di e en species in equilib ium and
o hei speci ic abili y o b eak he wine a oma bu e . Fu he mo e, pe cep ual
in e ac ions be ween he indi idual pe cep ions elici ed by each odo an will also
a ec he way hey a e pe cei ed by human senses (Fe ei a, 2010).
All he a oma-senso y a ibu es ound in wine a e caused by one o mo e
molecules which we e p esen in su icien concen a ion o su pass he wine
a oma bu e . The wine a oma bu e e e s o he speci ic senso y p ope ies o
he mix u e o 27 compounds om di e en chemical amilies ound in all wines
and alcoholic be e ages a he concen a ions p oduced in a no mal alcoholic
e men a ion. These molecules a e he main seconda y p oduc s o alcoholic
In oduc ion
10
e men a ion and a e esponsible o impo an p ocesses o a oma supp ession,
pa icula ly o ui y and woody no es (de-la-Fuen e-Blanco e al., 2016). The
composi ion o he wine bu e can sligh ly change since i depends on yeas
me abolism and o he oenological p ac ices, bu , all in all, i s senso y p o ile
does no change much and is desc ibed as “ inous” (Fe ei a e al., 2019).
An a oma ec o is de ined as “a pe cep ual uni cons i u ed by one o se e al
molecules wi h simila a oma desc ip o s, which al oge he and in an in eg a ed
o m, a e esponsible o a speci ic se o senso y ea u es o a ype o p oduc s;
wine in ou case” (Fe ei a e al., 2019).
Wine la ou is one o he mos complex and di icul o cha ac e ize and
manage, since he e is a huge a iabili y associa ed wi h i s o ma ion. Fac o s
like g ape a ie y, ine managemen , sani a y condi ions, loca ion and soil ype,
mic oo ganisms in ol ed in e men a ion, echnological choices o wine making
p ac ices, addi i es, wood ba els and wine p ese a ion a e only a ew examples
o impo an sou ces o a oma a iabili y (Fische e al., 1999; Robinson e al.,
2014).
1.1 Wine a oma genesis
Wine ola iles can be classi ied acco ding o hei chemical s uc u e, hei odou
in o a oma amilies, o acco ding o hei con ibu ion o a speci ic a oma ec o .
Howe e , in his wo k we a e mo e in e es ed in a classi ica ion acco ding o
hei genesis in o he ollowing ca ego ies:
1. A oma compounds de i ed om speci ic p ecu so s in he g ape
2. Fe men a i e a oma compounds om unspeci ic p ecu so s:
a. Rela ed o he g ape a ie y
b. Un ela ed o he g ape a ie y
3. A oma compounds o med o ex ac ed du ing aging
In oduc ion
11
The mos impo an g ape a ie ies o winemaking ha e a a he neu al a oma
cha ac e , howe e , hey con ain a se ies o speci ic a oma p ecu so s, which
a e a mo e o less complex chemical p ocess including hyd olysis, enzyma ic
clea age o spon aneous chemical ea angemen , ende he odo an .
Compounds in his ca ego y a e easily iden i ied as a ie al a oma compounds
because chemically hey ha e been buil by he g ape. Speci ic a oma p ecu so s
a e glycosidic p ecu so s and cys einyla ed and glu a hionyla ed p ecu so s
being he o me esponsible o he o ma ion o e penes, no isop enoids and
ola ile phenols and he la e o poly unc ional me cap ans (Fe ei a, 2010). A
hi d compound in his ca ego y is he amino acid S-me hyl me hionine, he
p ecu so o dime hyl sulphide (DMS) (Landaud e al., 2008).
Howe e , g apes also con ain a mo e o less speci ic p o ile o nu ien s which
will de e mine yeas me abolism and hence, also he e men a i e a oma p o ile.
Those g ape componen s in luencing yeas me abolism can be ega ded as
unspeci ic a oma p ecu so s ela ed o he g ape a ie y. These compounds a e
mos o en classi ied as e men a i e, since s uc u ally hey ha e been buil by
he yeas and no by he g ape, howe e , hey may ha e a e y impo an ole in
he iden i y o he speci ic a oma o he a ie y.
Impo an non-speci ic g ape p ecu so s a e amino acids which will lead o he
o ma ion o compounds like highe alcohols, b anched acids, hei e hyl es e s
and he ace a e es e s o highe alcohols. The speci ic p o ile o compounds o
hese chemical amilies o med du ing e men a ion is s ongly linked o he
speci ic g ape amino acid p o ile, howe e i s o ma ion occu s due o alcoholic
and/o malolac ic e men a ion (Fe ei a, 2010; He nández-O e e al., 2002;
Swiege s e al., 2005).
The e a e o cou se, o he e men a i e compounds o med om unspeci ic
a oma p ecu so s no ela ed wi h he g ape a ie y. E hanol is he mos
impo an in his ca ego y, bu hyd ogen sulphide (H2S), whose le els a e
In oduc ion
12
s ongly de e mined by he esidues o elemen al sulphu sp ayed o he ine,
can be also classi ied in his ca ego y (Ji anek e al., 1995; Mendes-Fe ei a e
al., 2009).
Finally, aged- ela ed a oma is o med by molecules ex ac ed om he wood
such as whiskylac ones, o o med by oxida ion o di e en p ecu so s, such as
s ecke aldehydes o so olon, o o med by he eac ion o wine componen s,
such as u u yl hiol which is o med by eac ion be ween H2S and u u al.
Li le amoun s o H2S o Me hane hiol (MeSH) o med by slow ca aly ical
decomposi ion o S-amino acids also belong o his ca ego y (Fe ei a, 2010;
Loscos e al., 2009).
1.2 G ape a ie y- ela ed non-speci ic p ecu so s
The main non-speci ic g ape p ecu so s a e g ape amino acids (Albe s e al.,
1996; He nández-O e e al., 2002). O he compounds which may be ega ded
as unspeci ic p ecu so s a e g ape lipids, no ably phy os e ols. A e y ecen
epo demons a es ha he wine a oma signa u e s ongly depends on he
p esence and ype o hese compounds (Fai bai n, 2018). Howe e , and o he
bes o ou knowledge, he e a e no u he clues abou he a oma compounds
ela ed o he p esence o hose compounds, al hough i can be hypo hesized ha
hey a e a y acids and hei co esponding es e s.
Amino acids a e he mos well-known g ape elemen s ela ed wi h odo an s.
Ea ly s udies e ealed ha le els o highe alcohols, hei ace a es, b anched
acids and hei e hyl es e s we e linked o he g ape a ie y wi h which he wine
was made (Fe ei a e al., 2000). La e , i was demons a ed ha he e men a ion
o syn he ic mus s con aining he cha ac e is ic amino acid p o iles o each
a ie y, e ec i ely p oduced speci ic a oma p o iles con aining all hese a oma
compounds (He nández-O e e al., 2002).
In oduc ion
13
1.3 Glycosidic p ecu so s
Glycosyla ion is a common anspo and de oxi ica ion plan mechanism and
hus has been desc ibed in se e al plan species (Sa y and Güna a, 2004;
Win e hal e and Skou oumounis, 1997). Glycosidic p ecu so s a e o med by
uni s con aining a β-D-glucose moie y (suga moie y) and an aglycone which will
p oduce he ola ile molecule (Win e hal e and Skou oumounis, 1997).
Common suga moie ies in g apes a e α-L-a abino u anosyl-β-D-
glucopy anoside, α-L- hamnopy anosyl-β-D-glucopy anoside, β-D-
xylopy anosyl-β-D-glucopy anoside, β-D-apio u anosyl-β-D-glucopy anoside,
and β-D-glucopy anoside-β-D-glucopy anoside. Rega ding he aglycone, he
di e si y is immense and includes e penes, C13-no isop enoids, C6-alcohols,
ola ile phenols and benzyl de i a i es (Liu e al., 2017; Wilson e al., 1984;
Win e hal e and Rouse , 2001; Win e hal e and Skou oumounis, 1997).
Glycosidic p ecu so s con ibu e o he a ie al exp ession o wine a oma and
a e a huge sou ce o a oma a iabili y, due o he high numbe o ac o s
a ec ing he composi ion o his ac ion and also o high numbe o ac o s
a ec ing he odo an s inally p oduced om i .
These p ecu so s we e ini ially iden i ied and s udied in he a oma ic a ie y
Musca liking mono e penes like linalool and ge aniol wi h i s a ie al cha ac e .
Thei impo ance was u he con i med in o he a ie ies like Gewü z amine
and Riesling (Güna a e al., 1985; S auss e al., 1986). O he ele an a oma
compounds like β-damascenone, cis- ose oxide, 1,1,6- ime hyl-1,2-
dyhyd onaph halene (TDN) o 4- inylphenol we e u he iden i ied in Riesling,
Gewü z amine and o he g ape a ie ies and we e also linked o glycosidic
p ecu so s (Pa ke e al., 2017; Se on e al., 2011; Win e hal e and Rouse ,
2001; Win e hal e and Skou oumounis, 1997).

In oduc ion
14
Some o hese p ecu so s a e able o p oduce a ee odo an by di ec hyd olysis
o he aglycone (S auss e al., 1986; Wilson e al., 1984), howe e o he
compounds can ha e mul iple in e media y p ecu so s, gene ally non- ola ile,
which by a se ies o ea angemen s will ul ima ely o igina e a ee ola ile. One
o he mos s udied cases a e he compounds de i ed om ca o enoids
(Win e hal e and Rouse , 2001).
Howe e , no mal glycosyla ion mechanism ha occu in g ape ines
(Win e hal e and Skou oumounis, 1997), a e no he only sou ce o glycosidic
p ecu so s. The plan is also able o glycosyla e ola ile molecules exis ing in
he ex e nal en i onmen being he bes -known case om smoke. The
appea ance o smoke- ain in wines a e la ge wild i es close o ineya ds was
a ibu ed o he glycosyla ion o ola ile phenols, such as guaiacol and 4-
me hylguaiacol, p esen in he a mosphe e. The s udy conduc ed by Kennison e
al., 2008 compa ing g ape juice and co esponding inal wine om ineya ds
exposed o smoke and unsmoked g ape ines, has shown ha ola ile phenols
p esen in smoke we e accumula ed as non- ola ile g ape glycosidic p ecu so s
in exposed g apes, bu hey we e only e ealed as ee odo an s in inal wine
a e mus e men a ion, eaching le els esul ing in consume ejec ion. Fu he
analysis on bo led aged wine showed ha le els con inued o inc ease indica ing
ha hese molecules kep on being hyd olysed om he glycosides by slow bu
spon aneous chemical hyd olysis.
1.4 Cys einyla ed and glu a hionyla ed p ecu so s
Cys einyla ed and glu a hionyla ed p ecu so s we e i s disco e ed in
Sau ignon Blanc and a e esponsible o he o ma ion o he po en a ie al
poly unc ional me cap ans in wine: 4-me cap o-4-me hyl-2-pen anone
(4M4M2P), 4-me cap o-4-me hyl-2-pen anol (4M4M2POH), 3-
In oduc ion
15
me cap ohexanol (3MH) and 3-me cap ohexyl ace a e (Da ie e al., 1993;
Peña-Gallego e al., 2012). These compounds a e p esen in wine in anges o
ng/L bu ha e an ex eme odo an po en ial o opical ui s, gua a, passion
ui , g ape ui o box ee. Depending on he concen a ion, hey can also be
pe cei ed as swea o onion (Fe ei a and San Juan, 2012; Ma eo-Vi a acho e
al., 2010; Peña-Gallego e al., 2012).
1.5 E ec s o ex e nal addi i es on wine a oma
In addi ion o g ape composi ion, also ex e nal addi i es can in luence he
o ma ion o a oma compounds du ing alcoholic e men a ion. These elemen s
include me al ca ions o en added as e ilize s o pes icides, SO2 added o
p ese e wine quali y o oak ba els o men ion only a ew examples. These
elemen s in eg a e wine p oduc ion p ocess and can highly a ec he
mic obiome ac i i y by modula ing hei enzyma ic esponse signal by he
p esence o ce ain me al ca ions (De Nicola e al., 2009) o by ha ing an isep ic
ac ion as in he case o SO2. Oak ba els can con ibu e highly o he ex ac ion
o wood molecules such as lac ones, howe e hey we e also linked wi h he
appea ance o o - la ou s by spoilage yeas ac i i y (Fe ei a, 2010; Mal ei o-
Fe ei a, 2011; Ribé eau-Gayon e al., 2006).
Zinc, o ins ance, is p esen in ineya ds and consequen ly in g ape mus . I has
a wide ange o sou ces om wa e pipes o pes icides (De Nicola e al., 2009;
Hop e e al., 2015). Yeas cells egula e he up ake o zinc by memb ane
anspo e s and he ac i a ion o inhibi ion o se e al yeas me abolic pa hways
a e egula ed by he zinc ex e nal concen a ion h ough a speci ic me al-
esponsi e egula o y p o ein (De Smid e al., 2008). This ca ion is c ucial o
he de elopmen o yeas s ains, since i in eg a es he ac i e-si e o 6 impo an
In oduc ion
16
classes o enzymes: oxido educ ases, ans e ases, hyd olases, lyases,
isome ases, and ligases (De Nicola e al., 2009).
SO2 is commonly used in winemaking due o i s an isep ic and an ioxidan
p ope ies. When added o he mus in su icien concen a ion, SO2 des oys
mos o wild spon aneous yeas and bac e ia, p e en ing spon aneous
e men a ion (Henick-Kling e al., 1998). La e on, du ing s o age, his
compound is also added o he wine o p e en ing he p oli e a ion o spoilage
mic oo ganism like ace ic acid bac e ia o B e anomyces spp., hus, a oiding
he o ma ion o o - la ou s like inega and ho se swea , espec i ely
(Mal ei o-Fe ei a, 2011; Ribé eau-Gayon e al., 2006).
Addi ionally, SO2 is highly eac i e wi h oxygen and ca bonyl compounds, hus
i p e en s di ec oxida ion by binding wi h dissol ed oxygen and by inhibi ing
oxida ion enzymes. On an indi ec manne , i p e en s ypical oxida ion la ou s
in wine by binding o aldehydes such as ace aldehyde (Fe ei a e al., 2015;
Ribé eau-Gayon e al., 2006).
2. Wine a oma o ma ion om speci ic and non-speci ic p ecu so s du ing
alcoholic e men a ion
Fe men a ions ha e a c ucial ole on he o ma ion o wine a oma. On one hand,
mic oo ganisms a e di ec ly esponsible o he o ma ion o many a oma
compounds ound ans e sely in all wines ( e men a i e compounds), bu hey
a e also impo an modula o s in he p oduc ion o a ie al la ou s om speci ic
p ecu so s. The o ma ion o hese las compounds is no , howe e , ully
unde s ood, despi e la ge esea ch conduc ed o da e.
In oduc ion
17
2.1 Spon aneous e sus inocula ion e men a ions
In na u e, se e al yeas gene a ha e been isola ed bo h om ineya ds and
wine ies, which demons a es he exis ence o a complex mic obiome in bo h
loca ions (Ba a a e al., 2012; Flee , 2003). S. ce e isiae has been desc ibed as
he “wine yeas ” due o i s high e men a i e igou and esis ance o o he
compe i o mic oo ganism and e en an isep ics like SO2 (Flee , 2003; Henick-
Kling e al., 1998; Jolly e al., 2014). Howe e , o he yeas gene a, mos ly non-
Saccha omyces yeas s, we e shown o be ac i e du ing he la en phase o no mal
alcoholic e men a ions a e which S. ce e isiae akes o e , ca ying mos pa
o he alcoholic e men a ion (Ba a a e al., 2012). Howe e , no all he yeas
ac i e we e conside ed posi i e and a he we e o en associa ed wi h he
o ma ion o o - la ou s like ace ic acid and e hyl ace a e (Jolly e al., 2014).
Fo his eason, o yea s, adi ional winemaking has p e e ed o use
comme cial d ied yeas p oduc s o isola ed S. ce e isiae o S. bayanus s ains.
The addi ion o a la ge numbe o cells o he g ape mus ensu es ha any
spon aneous mic oo ganisms a e su passed, ob aining a con olled e men a ion
and a oiding s uck o p oblema ic e men a ions.
This app oached ensu es indeed highe con ol o e e men a i e p ocesses, bu
has been demons a ed o lead o s anda diza ion o wines and hus, new s udies
ha e been done o assess which non-Saccha omyces gene a could lead o
posi i e ai s in wine (Henick-Kling e al., 1998; He nández-O e e al., 2008;
Jolly e al., 2014; Padilla e al., 2016).
Du ing alcoholic e men a ion, S. ce e isiae is no only esponsible o he
con e sion o glucose and uc ose in o e hanol and CO2, bu also o he
o ma ion o impo an ola iles due o he seconda y yeas me abolism. The
wine me abolome is dependen on he pool o g ape p ecu so s and on he
e iciency wi h which hey a e con e ed in o ola ile molecules du ing
alcoholic e men a ion. The o ma ion o ola ile compounds h ough yeas
In oduc ion
24
Bu , acid ca alysis is also esponsible o he ans o ma ion o ele an a oma
ola iles, such as ge aniol, which a e being eleased om he glycoside, will
spon aneously ans o m in o he much less odo an a- e pineol, as shown in
Figu e 4.
2.3.2 Release and o ma ion o no isop enoids
The mos impo an a oma compounds de i ed om no isop enoids a e β-
damascenone, β-ionone and TDN. Some o he a oma molecules wi h less
a oma ic ele ance a e a-ionone, Riesling ace al and i ispi anes. All hese
molecules a e chemically ke ones, hyd oca bons o e he s, meaning ha hey
canno be pa o aglycones, which by na u e ha e o ha e a hyd oxyl g oup
(alcohols o ca boxylic acids). This implies ha he o ma ion o hese a oma
Figu e 4 Fo ma ion o mono e penes om glycosidic p ecu so s due o enzyma ic o acidic hyd olysis (a)
and (b) ollowed by chemical ea angemen s o o m o he mono e pene molecules due o acidic
en i onmen (a) adap ed om Wa e house e al., 2016.

In oduc ion
25
molecules will, necessa ily, ake place a e di e en eac ions o he han he
clea age o he glycosidic p ecu so s. Fo ins ance, one possible mechanism o
β-damascenone o ma ion in wine has been shown o be om he ca o enoid
neoxan hin, equi ing oxida i e clea age, ollowed by enzyma ic educ ion and
inally acid hyd olysis in o de o become ola ile (Win e hal e and Rouse ,
2001).
While all no isop enoids de i e om ca o enoid b eakdown, wo possible
o ma ion ou es a e possible ( igu e 5):
1. he di ec deg ada ion om ca o enoids p esen in he g ape mus o in
he wine, as seems o be he case o β-ionone (Win e hal e and Rouse ,
2001)
2. he clea age and u he ea angemen o non- ola ile glycosides o C13-
no isop enoids in e media es o med om ca o enoids (β-damascenone,
TDN, i ispi ane o Riesling ace al (Win e hal e , 1991; Win e hal e
and Rouse , 2001).
Figu e 5 Possible pa hways o o m ola ile no isop enoids compounds om ca o enoid ini ial p ecu so s
in ol ing mul iple glycosyla ed in e media es and/o chemical, enzyma ic o acid hyd olysis In Mendes-
Pin o, 2009.
In oduc ion
26
One o he di icul ies is ha o all he impo an odo an s, he e a e mul iple
p ecu so s, whose s uc u es a e no well known. This was i s obse ed by
Pe e Win e hal e in 1991 o TDN, i ispi ane and Riesling ace al, bu seems
o be alid also o β-damascenone. To complica e mo e hings, he same
p ecu so s can yield di e en molecules, and some o hem, such as Riesling
ace al, can be in e media es in he p oduc ion o TDN (Gök, 2015).
2.3.3 Release and o ma ion o ola ile phenols
Vola ile phenols such as guaiacol, anillin, c esols and eugenol can be ex ac ed
om wood (oak ba els, wood chips, e c) con ibu ing wi h smoky, swee o
clo e la ou s o wine (Kennison e al., 2008). Small amoun s o hese ola iles
a e also p esen unde he o m o glycosides in g apes, so ha hese compounds
can be also eleased by enzyma ic o acid hyd olysis.
Mo eo e , g apes con ain a la ge a ie y o phenolic compounds including
phenolic acids, la onoids, an hocyanins o annins (Khei e al., 2013). Majo
di e ences exis be ween whi e and g ape a ie ies, which p ecisely p o ides
colou di e ences in wine. Howe e , some o hese phenolic compounds a e
ac ual a oma p ecu so s o ola ile phenols, such as inyl phenols, which can be
also ound as glycosidic p ecu so s. Vinyl phenols a e mo e impo an in whi e
wines since he biochemical eac ions o he ans o ma ion o phenolic acids
a e inhibi ed by ed wine componen s (Basha e al., 2004; Cha onne e al.,
1993).
Phenolic acids in g ape a e di ided in o wo g oups benzoic and cinnamic acids
and a e p esen in la ge amoun in ed g apes. Cinnamic acids exis in g ape
be ies unde di e en o ms, since in addi ion o he ee molecules, hey can
also be as es e s o a a ic acid and as glycosides (Basha e al., 2004; Khei e
al., 2013). S. ce e isiae is able o o m inylphenols du ing alcoholic
In oduc ion
27
e men a ion om cinnamic acids, in pa icula e ulic and p-couma ic acid
ca alysed by he enzyme cinnama e deca boxylase (Cha onne e al., 1993).
The con e sion o inylphenols in o e hyl phenols is a e in mic oo ganisms and
la ge amoun s o hese compounds ha e been linked wi h spoilage yeas
B e anomyces spp. (Khei e al., 2013).
3. Selec ion o non-Saccha omyces yeas s ains
Non-Saccha omyces yeas s a e a ely used as single inoculums since hey a e
inhibi ed by high e hanol le els and hei suga me abolism is no as e icien as
ha o S. ce e isiae. In ac , non-Saccha omyces yeas s ha e been in he pas
associa ed wi h sluggish o s uck e men a ions wi h an inc eased isk o
spoilage by compe i o mic oo ganisms such as ace ic acid bac e ia (Flee ,
2003). These non-Saccha omyces a e hen used as mixed inoculum o ollowing
sequen ial inocula ion p ocedu es in which he inal pa o he e men a ion is
ca ied ou by S. ce e isiae.
In bo h cases, non-Saccha omyces yeas s can ac as po en ial enhance s o wine
a oma due o hei po en ially di e en enzyma ic ac i i y compa ed o S.
ce e isiae, namely by hei speci ic b-glucosidase and b-lyase ac i i ies
(Mendes Fe ei a e al., 2001; Zo e al., 2011), bu he e a e many o he a oma
compounds whose le els a e modula ed by he p esence o hese yeas s. The
modula ion may ake ia di e en mechanisms no ye clea , such a di ec
in e ac ion be ween yeas s ains o compe i ion. The educ ion o nu ien s, he
ea ly amino acid in ake o e en limi a ions in dissol ed oxygen could be
impo an modula o s o yeas me abolism (Clemen e-Jimenez e al., 2005;
Flee , 2003; Kapsopoulou e al., 2007; Mo eno e al., 1991). On he o he hand,
some s ains o non-Saccha omyces wi h abili ies o sec e e lipoly ic enzymes o
In oduc ion
28
he media, may deg ade g ape lipids in o ee a y acids, compounds which can
inhibi he g ow h o S. ce e isiae (Esc ibano e al., 2017).
Th ee comme cial s ains, isola ed om wine en i onmen s, a e o en used in
wine making and ha e shown pa icula ai s.
3.1 To ulaspo a delb ueckii
T. delb ueckii was one o he i s non-Saccha omyces comme cial s ains. I is
desc ibed as ha ing medium e men a i e powe and o be a low p oduce o
ace ic acid. When co-inocula ed wi h S. ce e isiae, i was shown o imp o e he
senso y desc ip ion o wine, howe e i is no clea how his was accomplished,
since esea che s desc ibed ha co-inocula ion induced he dec ease o isoamyl
ace a e, o a y acids wi h known ole on ui y a omas, such as hexanoic acid
and also o inyl phenols (Azzolini e al., 2014; Jolly e al., 2014). Resea che s
ha e also sugges ed ha his s ain has ele an β-glycosidase and ca bon-sul u
lyase ac i i ies (Esc ibano e al., 2017; Padilla e al., 2016).
3.2 Pichia kluy e i
P. kluy e i was i s associa ed wi h a highe elease o ola ile hiols (An ang
e al., 2009), bu i has been also linked wi h he o ma ion o ace a e es e s
(Viana e al., 2008). Compa ing o o he yeas s ains, his yeas has a qui e
speci ic me abolic cha ac e is ics including an oxida i e me abolism
cha ac e ized by he o ma ion o bio ilms (Ba a a e al., 2012).
3.3 Lachancea he mo ole ans
This yeas was i s desc ibed by i s abili y o p oduce lac ic acid and hus,
con ibu e o he wine oundness (Kapsopoulou e al., 2007; Va ela and
Bo neman, 2017). Besides, senso y desc ip ou s such as spiciness whe e ound
o inc ease in wines e men ed wi h his yeas (Gobbi e al., 2013).

Bibliog aphy
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Food Chemis y 55, 6674–6684.

Sec ion I
S ecke aldehydes a e no mal by-p oduc s o
alcoholic e men a ion linked o yeas sulphi e
me abolism
Sec ion I - In oduc ion
43
1. In oduc ion
S ecke aldehydes (SAs), namely 2-me hylp opanal, 3-me hylbu anal, 2-
me hylbu anal, me hional and phenylace aldehyde, a e powe ul a oma
molecules playing ele an oles in he la ou o wine and bee . In bee , 3-
me hylbu anal and 2-me hylbu anal we e i s p oposed as esponsible o he
mal la ou , no e cha ac e is ic o some alcohol- ee bee s (Beal and Mo am,
1994). O he au ho s la e demons a ed ha me hional was in ac mo e ele an
in such o -odou (Pe pe e and Collin, 1999a). Fu he mo e, he implica ion o
me hional and phenylace aldehyde in some nega i e odou cha ac e is ics o
oxidized/aged bee was demons a ed in 2004 (Soa es da Cos a e al., 2004) and
con i med in mo e ecen s udies (Saison e al., 2010; Wie s ock e al., 2016). In
he case o wine, he in ol emen o me hional in he “cooked ege ables” no e
o oxidized wines was s ablished in 2000 (Escude o e al., 2000) and
phenylace aldehyde was iden i ied by GC-O (GC-Ol ac ome y) as one o he
key odo an s o oxidized whi e wines in 2003 (Sil a Fe ei a e al., 2003). The
main oles played by hese compounds in he odou no es o oxidized wines we e
u he con i med in 2007 (Culle e e al., 2007).
These ubiqui ous and powe ul smelling molecules a e chemically o
biochemically ela ed o he so called S ecke amino acids: aline, leucine,
isoleucine, me hionine and phenylalanine. The oxida i e deamina ion o hese
amino acids in he p esence o ea polyphenols o o m he co esponding
aldehydes was obse ed as soon as 1954 and was con i med in he 70’s when
Japanese esea che s demons a ed ha he deg ada ion in ol ed he eac ion o
he amino acid wi h a quinone de i ed om a la anol unde going oxida ion
(Saijō and Takeo, 1970a). The chemical ou es leading o he o ma ion o hese
aldehydes in oxida ion- ela ed p ocesses ha e been ela i ely well es ablished
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
44
(Bae e al., 2012; Bueno e al., 2018; G an -P eece e al., 2013; Rizzi, 2006;
Wie s ock e al., 2016).
The e men a i e o igin o hese compounds was i s sugges ed also by (Saijō
and Takeo, 1970b) who obse ed ha supplemen ing e men ing ea lea es wi h
phenylalanine esul ed in phenylace aldehyde o ma ion. Nowadays, i is known
ha SAs a e no mal in e media es o he Eh lich pa hway in he amino acid
me abolism, bu he cu en belie es es ablish ha hey a e mos ly educed o he
co esponding alcohols, so ha le els o SAs in esh bee o wines a e hough
o be negligible. Howe e , he e a e some e idences poin ing ou ha in ce ain
condi ions, yeas canno educe all he SAs. This was i s obse ed in cold
e men a ion condi ions o he p oduc ion o alcohol- ee bee (Pe pe e and
Collin, 2000a). Such inabili y was en a i ely a ibu ed o he p esence o
sulphi e o la onoids (Pe pe e and Collin, 2000b). O he esea che s u he
con i med ha e e men a ion o aged bee educed bu was no able o
comple ely elimina e SAs and ha esidual le els we e s ain dependen (Saison
e al., 2010). I.e., hese wo ks show ha esea che s ha e been long awa e o he
ac ha aldehydes can o m s able and e e sible non- ola ile adduc s wi h SO2
(Bae e al., 2012; de Aze edo e al., 2007), ha such adduc s could limi he
e iciency o yeas educ ases, and ha adduc s could also play some ole in he
ul e io de elopmen o oxidized no es. In spi e o his e idence, he po en ial
impo ance o alcoholic e men a ion as a ele an sou ce o SAs emains
unexplo ed.
Recen ly, we de eloped an analy ical me hod able o measu e ee aldehydes and
o es ima e he bonded ac ion (Bueno e al., 2014). Using such me hodology,
i was possible o con i m ha non-oxidized wines may con ain a la ge pool o
SAs unde he o m o sulphi e adduc s. These adduc s a e p og essi ely clea ed
Sec ion I - In oduc ion
45
du ing he i s s ages o wine oxida ion as ee SO2 is deple ed, concomi an ly
eleasing he ee o ms o he aldehydes (Bueno e al., 2016). The p oduc ion o
SAs om oxida i e deg ada ion o amino acids was ound o ake place only
when le els o ee SO2 become smalle han 4 mg/L, sugges ing ha oxida i e
odou no es de eloped by some wines du ing aging could be in ac due o he
simple elease o sulphi e adduc s, and no o he oxida ion o amino acids,
alcohols o o he p ecu so s. Fu he mo e, PLS modelling sugges ed ha SAs
p esen in no mal non-oxidized wines as sulphi e adduc s, could ha e been
o med in e men a ion as a consequence o a ailu e in he ac ion o alcohol
dehyd ogenases, possibly induced by a lack o zinc, and likely by he aldehyde-
p o ec ing ac ion o SO2 (Bueno e al., 2016).
In o de o con i m hose e idences, he esea ch p esen ed in his pape , s udies
he o ma ion o SAs in alcoholic e men a ion using syn he ic media esembling
g ape mus . The majo objec i es a e o assess he e ec s o he s ain o yeas
and o he le els o zinc and SO2 o he ini ial mus on he le els o SAs o med
in he alcoholic e men a ion.

S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
46
2. Me hodology
2.1 Reagen s and s anda ds
Sodium me abisul i e (97%), zinc chlo ide (97%), hyd ogen pe oxide 3%
s abilized w/ , indica o 4.4 mixed (me hyl ed-me hylene blue), sodium
hyd oxide 0.01 mol/L, o ho phospho ic acid (85%) o VINIKIT line we e
ob ained om Pan eac (Ba celona, Spain).
Dichlo ome hane (DCM), e hanol and me hanol (≥ 99%) wi h Dis ol-Pes icide
esidue g ade we e supplied by Me ck (Da ms ad , Ge many). Glyoxal solu ion
40 w . % in H2O was pu chased om Sigma-Ald ich (Mad id, Spain).
The in e nal s anda ds me hyl 2-me hylbu y a e (≥ 99%) and 2-bu anol (≥ 99%)
we e ob ained om Me ck, while 4-me hyl-2-pen anol (99%), 4-hyd oxi-4-
me hyl-2-pen anone (99%), e hyl hep anoa e (99%) 2-oc anol (99.5%) and
hep anoic acid (99 %) we e pu chased om Sigma Ald ich. Wa e was pu i ied
using Milli-Q® sys em om Millipo e (Me ck).
2.2 Cul u e condi ions:
2.2.1 Syn he ic mus composi ion
Syn he ic mus : he syn he ic mus esembling g ape mus was adap ed om
Bely, Sablay olles, & Ba e, 1990 and had he ollowing composi ion:
Oligoelemen s: MnCl2.4H2O 4.7 mg/L, Co(NO3)2·6H2O 0.49 mg/L,
NaMoO4·2H2O 0.19 mg/L, CuCl2 0.54 mg/L, KIO3 1.29 mg/L, H3BO3 1 mg/L,
SO4Mg·7H2O 0.2 g/L, KH2PO4 2 g/L, CaCl2·2H2O 0.155 g/L; Acids: malic acid
0.3 g/L, a a ic acid 3 g/L, ci ic acid 0.3 g/L, wi h pH adjus ed o 3.5 wi h HCl;
Vi amins -all supplied om Me ck (≥ 98%): py idoxine hyd ochlo ide 1 mg/L,
nico inic acid 1 mg/L, calcium pan o hena e 1 mg/L, hiamine hyd ochlo ide 1
mg/L, p-aminobenzoic acid 1 mg/L, ibo la in 0.2 mg/L, olic acid 0.2 mg/L,
Sec ion I - Me hodology
47
bio in 0.04 mg/L; myo-inosi ol (≥ 99%) 0.3 g/L, e gos e ol (≥ 75%) 15 mg/L;
Suga s we e om Pan eac Applichem (Spain): glucose 100 g/L; uc ose 100
g/L; ween 80® 0.05 % ( / ) (Sigma-Ald ich); ni ogen sou ce: (NH4)2HPO4
0.2199 g/L; amino acids (Me ck) (mg/L): GABA 44.37, alanine 58.51, y osine
14.34, aline 17.73, isoleucine 14.43, leucine 13.42, aspa a e 34.82, glu amic
acid 61.83, glu amine 104.83, se ine 21.21, glycine 1.11, his idine 109.2,
h eonine 18.8, a ginine 199.5, p oline 241.46, me hionine 29.85, phenylalanine
11.15, lysine 3.33.
Zinc was added om a s ock solu ion o ZnCl2, 162 mg/L.
SO2 was added om a eshly p epa ed Na2S2O5 solu ion, 5000 mg/L.
2.2.2 Yeas cul u e and e men a ion se -up
Yeas cul u e: h ee comme cial Saccha omyces ce e isiae yeas s ains we e
selec ed: L1 - Q23 (Lallemand), L2 - Me i (Ch . Hansen); L3 - Fe mic u AR2
(Oenob ands). The yeas cells we e hyd a ed o 1 hou a 35°C ollowed by he
addi ion o syn he ic mus o ac i a e hem a he same empe a u e o a ound
30 min o 1hou . The e men e s we e inocula ed wi h 106 cells/ml.
Mus manipula ion: he syn he ic mus was s e ilized by means o s e ile
cellulose ni a e memb ane il e s wi h 0.45 µm po es (Albe , A&S Fil e Co.,
L d) a e i s p epa a ion and pH adjus men . All media manipula ions we e
made inside a e ical lamina low chambe PV-100 (Tels a , S.A), o ensu e
wo king unde asep ic condi ions. Small olumes we e s e ilized using sy inge
il e s wi h 0.2 µm HT Tu yn® memb ane om PALL (New Yo k, USA). All
glasswa e was s e ilized using an au ocla e AES-28 om Raypa (Ba celona,
Spain).
Fe men a ion se -up: The i s expe imen al se -up was pe o med wi hou SO2
and he concen a ions o zinc es ed we e 10 mg/L, 5 mg/L and 1 mg/L. In he
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
48
second se -up all samples con ained 30 mg/L o SO2 and he concen a ions o
zinc we e 10 mg/L, 1 mg/L and 0 mg/L. Two biological eplica es con aining
150 mL o he mus we e used o each condi ion.
Fe men a ion: Fe men a ions we e ca ied ou in 250 mL blue cap glass lasks
(Ilmabo TGI, Ge many) closed wi h ai lock al es and we e kep a cons an
empe a u e o 20°C. The p og ess o e men a ion was moni o ed by daily
con ol o he weigh . Fe men a ion was conside ed inished when he loss o
weigh be ween wo consecu i e days was smalle han 0.1 g.
Once e men a ion was conside ed inished, he e men e s we e sealed and
sonica ed o 15 minu es and we e hen in oduced in an anoxic chambe om
Jacomex (Dagneux, F ance) whe e hey we e le o sedimen o 5 hou s and
we e hen aliquo ed. Aldehyde and SO2 analysis we e pe o med on he
ollowing hou s, while he aliquo s o he emaining a oma compounds we e
p ese ed in ials in he idge.
2.3 Analy ical me hods
2.3.1 Classical oenological cha ac e iza ion
The wines we e cha ac e ized acco ding o hei gene al enological pa ame e s
using he ecommended me hodologies by OIV (In e na ional O ganiza ion o
Vine and Wine): educing suga s, o al acidi y and ola ile acidi y (In e na ional
O ganisa ion o ine and wine, 2011).
pH was measu ed using a pHme e . Wine o al acidi y was measu ed by i a ion
wi h NaOH 0.1 N. Vola ile acidi y was i a ion wi h NaOH 0.02 M o he
ola ile ac ion ob ained by s eam dis illa ion. Residual suga s we e calcula ed
using he Fehling p ocedu e based on oxida ion o educing suga s wi h CuII in
alkaline media by boiling he solu ion. Excess o cuppe no oxidized by he
Sec ion I - Me hodology
49
suga s, oxidases iodine added as KI. The solu ion is hen i a ed wi h sodium
hiosul a e using s a ch as indica o o de e mine I2.
F ee sulphu dioxide was de e mined by HeadSpace Gas Ch oma og aphy Mass
Spec ome y (HS-GC-MS) using a GCMS-QP2010 om Shimadzu (Kyo o,
Japan) as desc ibed in he li e a u e (Ca ascón e al., 2017). A DB-WAX
column was used (30 m x 0.25 mm i.d x 0.25 µm ilm hickness) om J&W
Scien i ic (Agilen Technologies, San a Cla a, CA USA). This me hod is based
on he displacemen o SO2 equilib ium o ms wi h o hophospho ic acid (85%),
in which 4.5 mL o wine wi h 20 µL o 2-clo oe hanol (in e nal s anda d) a e
ca ully capped in a 10 mL headspace ial. Jus be o e he analysis he sample is
acidi ied wi h 500 µL o o hophospho ic acid (85%).
To al SO2 was analysed using he aspi a ion/ i a ion me hod desc ibed by
Rankine and ecommended by he OIV (In e na ional O ganisa ion o ine and
wine, 2011). 3 mL o he hyd ogen pe oxide 3% (p/ ) wi h 3 d ops o me hyl
ed-me hylene blue indica o and 2-3 d ops o NaOH 0.01 M, so ha he solu ion
u ns om pu ple o g een, a e p epa ed in a hea -shape lask wi h a bubble .
The 10 mL o wine a e ans e ed o a ound lask wi h 5 mL o H3PO4 a 20%
a e secu ed in he wa e acuum sys em wi h a bubble ube. The ound lask is
hea ed and he aspi a ion sys em is ac i a ed o 15 minu es. To al SO2 is
measu ed by i a ion wi h NaOH 0.01 M.
2.3.2 Quan i ica ion o o al aldehydes
To al aldehydes we e analysed using a p e iously desc ibed me hod (Bueno e
al., 2014). Aldehyde-sulphi e adduc s (hyd oxyalkylsul ona es) we e p e iously
clea ed by incuba ing he wine in s ic anoxic condi ions wi h 6 g/L o glyoxal
a 50°C du ing 6 hou s. Released aldehydes a e u he analysed by HeadSpace
Solid Phase Mic oEx ac ion Gas Ch oma og aphy Mass Spec ome y (HS-
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
56
added inal
yeas zinc yeas *zinc yeas zinc yeas *zinc SO2SO2
o al SO20.000 0.033 - 0.000 - 0.040 0.000 0.002
aldehydes
2-me hylp opanal - - - 0.003 - - - -
3-me hylbu anal - - - 0.011 0.002 0.011 0.018 0.000
2-me hylbu anal - - - 0.003 - - - -
me hional 0.000 - - - - - 0.01 -
phenylace aldehyde 0.007 - - 0.000 0.001 0.000 0.000 0.000
usel alcohols
isobu anol 0.000 - - 0.000 0.014 0.047 0.006 0.000
isoamyl alcohol 0.000 - - 0.000 0.025 - 0.001 0.000
me hionol 0.001 - - 0.000 - - - -
2-phenyle hanol 0.002 0.010 - 0.000 0.010 0.009 0.000 -
iso-acids
2-me hylp opanoic acid 0.000 - - 0.000 - - - 0.000
3-me hylbu anoic acid 0.000 0.010 - 0.000 - - - 0.000
aldehyde/alcohol a io
2-me hylp opanal/ isobu anol - - - 0.003 - - - 0.000
3-me hylbu anal/ isoamyl
alcohol
0.019 - - 0.001 0.007 0.041 - 0.000
me hional/ me hionol 0.002 - - 0.001 0.021 - 0.002 -
phenylace aldehyde / 2-
phenyle hanol
0.001 - - 0.000 0.000 0.000 0.000 0.000
aldehyde/acid a io
2-me hylp opanal/ 2-
me hylp opanoic acid
- - - 0.049 - - - 0.000
3-me hylbu anal/ 3-
me hylbu anoic acid
- - - 0.000 0.002 0.003 - 0.000
isobu anol/2-me hylp opanoic
acid
0.000 - - - 0.042 - 0.006 -
isoamyl alcohol/ 3-
me hylbu anoic acid
0.008 - - - 0.043 - - -
2-me hylbu anal/
iso ale aldeyde
0.000 - 0.046 0.015 - - 0.004 0.003
medium chain a y acids
hexanoic acid 0.000 - 0.019 0.000 0.008 - 0.019 -
oc anoic acid 0.000 0.027 - 0.000 - - - -
decanoic acid 0.001 - 0.029 0.009 - - - -
es e s
isoamyl ace a e 0.000 - - 0.000 - - 0.000 -
e hyl hexanoa e - 0.000 - - 0.000 - 0.004 -
e hyl oc anoa e 0.001 0.000 0.006 0.023 0.000 - - -
e hyl decanoa e 0.000 0.000 0.001 0.032 0.001 - 0.001 -
es e /acid a ios
e hyl hexanoa e/ hexanoic acid 0.010 0.000 - - 0.000 - 0.000 -
e hyl oc anoa e/ oc anoic acid - 0.000 - - 0.000 - - -
e hyl decanoa e/ decanoic acid - 0.000 - - 0.000 - 0.000 -
2-way ANOVA
co ela ion
wi hou SO2
wi h SO2
Table 1 Summa y o he signi icance o he e ec s played by he ac o s yeas and zinc as well as hei
in e ac ion on he le els o o al SO2, SAs and majo e men a ion ola iles assessed by wo way-ANOVAs
ca ied ou in he wo da a se s; addi ion o ex e nal SO2 and he le els o o al SO2 ound a e
e men a ion gi en by signi icance o hei co ela ion wi h compounds and a ios.

Sec ion I – Resul s and Discussion
57
The abili y o S. ce e isiae o o m and exc e e SAs du ing cold con ac
e men a ion in he p oduc ion o low alcoholic bee s was demons a ed ime ago
(Pe pe e and Collin, 2000a). Simila conclusions we e eached s udying he
educ ion o aldehydes by e- e men a ion (Saison e al., 2010). I is, howe e ,
belie ed ha he o ma ion o aldehydes by e men ing yeas in bee is mos
likely limi ed and o sca ce impo ance, a leas in compa ison wi h o he sou ces
linked o wo p oduc ion (Bae e al., 2012). The expe imen s ca ied ou by
Saison e al., 2010, much in acco dance wi h all obse a ions ega ding he
abili y o yeas o educe aldehydes du ing e men a ion (Peppa d and Halsey,
1981), sugges ha SAs p oduced du ing mal ing and boiling will be educed o
he co esponding alcohols du ing e men a ion, bu ha he e is a ac ion o
SAs which will emain and is dependen on he yeas s ain and wo
composi ion. Likewise, his wo k he eby con i ms ha , du ing alcoholic
e men a ion o g ape mus , a ac ion o SAs a senso y ele an le els emains
in he wine and his is highly depend on he yeas s ain, as seen in Table 1.
Rega ding he pa icula e ec s o each s ain on le els o SAs, da a in Table 2
e eal ha le els o med a e speci ic o each case. L1 p oduces in gene al
smalle le els, eaching smalles alues o 2-me hylp opanal and 2-
me hylbu anal, bu le els o phenylace aldehyde p oduced by his s ain we e
ela i ely la ge. On he o he hand, L3 p oduced maxima alues o 3-
me hylbu anal and phenylace aldehyde, bu le els o me hional we e close o
hose p oduced by L1. In con as , L2 p oduced maxima le els o me hional and
minima o phenylace aldehyde. This complex pa e n o dependence would ha e
been expec ed, since hese aldehydes a e p oduced du ing he syn hesis o amino
acids and a e u he educed o alcohols by a complex and he e ogeneous
enzyma ic sys em which has o es o e he cell edox cycle (Peppa d and Halsey,
1981). This pool is in eg a ed by alcohol dehyd ogenases (ADHs), aldehyde
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
58
dehyd ogenases and aldoke o educ ases using ei he NAD(H) o NADP(H) as
co ac o s (Pe pe e and Collin, 1999b; Van Ie sel e al., 1997).
A ending o he p e ious discussion, le els o SAs e ained in he wines a e
e men a ion, should hen be ela ed o he amoun o highe alcohols p oduced
by he s ain, o he selec i i y and e ec i i y o he ADH-sys em o he s ain,
and e en ually, o he di e en ial le el o any molecula species able o p o ec
he aldehyde om educ ion o oxida ion, such as SO2 (Pe pe e and Collin,
2000b). The ac ha he a ios be ween he le els o aldehyde and hose o he
co esponding alcohol a e also signi ican ly ela ed o he yeas s ain (Tables 1
and 2), suppo s ha he speci ic abili y o each s ain o educe hese aldehydes
exe s a majo ole on he inal le el o emaining aldehydes. Wines made wi h
L3 ha e maxima aldehyde/ alcohol a ios (excep me hional/ me hionol) and
e y low le els o highe alcohols (excep me hionol). On he con a y, wines
made wi h L2 con ained maxima le els o highe alcohols (excep 2-
phenyle hanol) and lowes aldehyde/alcohol a ios (excep
me hional/me hionol), sugges ing ha his s ain educes aldehydes o alcohols
e icien ly.
Sec ion I – Resul s and Discussion
59
L1 L2 L3
o al SO2*22.1 ± 2b11 ± 1a32.8 ± 3c
aldehydes
2-me hylp opanal
3.6 ± 0.4a7.2 ± 0.7b6.1 ± 0.8b
3-me hylbu anal*23.8 ± 2.1a23 ± 1.8 a 30.4 ± 3.2b
2-me hylbu anal
1.7 ± 0.1 a 2.6 ± 0.2b2.7 ± 0.3b
me hional
23.7 ± 0.6 a 28.8 ± 0.7b24.7 ± 1.4a
phenylace aldehyde*14.2 ± 2.9b8.2 ± 1.5a18.6 ± 4.3c
usel alcohols
isobu anol*13.9 ± 0.5a41.1 ± 2.3c18.1 ± 0.9b
isoamyl alcohol
163 ± 6b242 ± 9c135 ± 10a
me hionol
4 ± 0.2 a 6.1 ± 0.2b6.1 ± 0.3b
2-phenyle hanol*27.1 ±1.5c23 ± 0.9b20 ± 1.5a
iso-acids
2-me hylp opanoic acid
0.97 ± 0.0a2.2 ± 0.1c1.2 ± 0.1b
3-me hylbu anoic acid
1.2 ± 0.0a1.9 ± 0.1b1.2 ± 0.1a
aldehyde/alcohol a io
isobu i aldehyde/ isobu anol
0.26 ± 0.0ab 0.2 ± 0.0a0.4 ± 0.0c
3-me hylbu anal/ isoamyl alcohol*0.15 ± 0.01a0.1 ± 0.0a0.2 ± 0.0 b
me hional/ me hionol
6.3 ± 0.5c4.8 ± 0.2b4.2 ± 0.2a
phenylace aldehyde/ 2-phenyle hanol*0.5 ± 0.1a0.4 ± 0.1a0.88 b
2-me hylp opanal/ 2-me hylp opanoic acid - - -
3-me hylbu anal/ 3-me hylbu anoic acid*20.6 ± 1.8 b 12.3 ± 1.1a26.5 ± 3.8c
isobu anol/ 2-me hylp opanoic acid
14.8 ± 1a18.9 ± 1b15.4 ± 1.1a
isoamyl alcohol/ 3-me hylbu anoic acid
141 ± 7b129 ± 6ab 112.5 ± 8.5a
2-me hylbu anal/ 3-me hylbu anal•0.1 ± 0.0a0.1 ± 0.0b0.1 ± 0.0 a
medium chain a y acids
hexanoic acid•1.9 ± 0.0c1.2± 0.0b1 ± 0.1a
oc anoic acid
3.7 ± 0.2b2.4 ± 0.1a2.3 ± 0.1a
decanoic acid•0.9 ± 0.1c0.6 ± 0.0 b 0.4 ± 0.0a
es e s
isoamyl ace a e
0.3 ± 0.0b0.5 ± 0.0c0.2 ± 0.0a
e hyl hexanoa e - - -
e hyl oc anoa e•3.7 ± 0.2b2.4 ± 0.1a2.3 ± 0.1a
e hyl decanoa e•0.3 ± 0.1b0.2 ± 0.0b0.1 ± 0.0a
a ios
e hyl hexanoa e/ hexanoic acid
14.5 ± 2.4a23 ± 5.9b23.6 ± 5.4b
e hyl oc anoa e/ oc anoic acid - - -
e hyl decanoa e/ decanoic acid - - -
Table 2 A e age le els o o al SO2, a oma compounds and o some ele an a ios a ending o he yeas
s ain. Concen a ion da a a e in mg/L excep aldehydes which a e in
µ
g/L. Cases showing signi ican
in e ac ions yeas x zinc a e ma ked wi h • o * o he expe imen s wi hou o wi h SO2, espec i ely.
Signi ican di e ences a ending o Duncan es a e indica ed wi h le e s a-c being “a” he lowes a e age
alue.
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
60
3.2 Role o SO2
I is wo h men ioning ha he inal le els o o al SO2 ound in he e men ing
media we e in ac mo e ela ed wi h he yeas s ain han wi h he ini ial le el
o SO2 added o he mus , as can be seen in Table 2 and in Figu e 7. Table 2
shows ha he a e age le els o SO2 emaining in samples e men ed wi h L3
we e he highes , wi h 32.8 mg/L, whe eas hose emaining in samples e men ed
wi h L2 we e he lowes , wi h 11.03 mg/L. Figu e 7 u he illus a es ha all
samples e men ed wi h L2 had inal le els o o al SO2 below 15 mg/L, e en i
he ini ial mus con ained 30 mg/L o his an ioxidan . On he o he hand,
samples e men ed wi h L3 wi hou ex e nal SO2 con ained 20-25 mg/L a he
end o e men a ion, and he ex e nal addi ion b ough abou an ex a inc ease o
nea ly 20 mg/L. I is ob ious ha each yeas s ain me abolizes SO2 di e en ly,
using i ei he as sou ce o sulphu o , on he con a y, p oducing i om o he
sulphu sou ces, mos likely o ake ad an age o i s oxic e ec on compe ing
mic oo ganisms. In ac , he esis ance o SO2 is a gene ically de e mined
cha ac e is ic o yeas s linked wi h an in e es ing molecula mechanism only
obse ed in wine s ains (Pe ez-O in e al., 2002) which has ecei ed some
a en ion o i s po en ial indus ial in e es (Di ol e al., 2012). Mo eo e , Nadai
e al., 2016 ha e ecen ly ound ha s ains showing highe esis ance o SO2,
p oduced highe le els o SO2 in compa ison o hose sensi i e o his molecule,
which sugges s ha SO2 esis ance and p oduc ion a e ela ed. Fu he mo e,
esis an s ains we e shown o ha e much highe basal gene exp ession le el o
SSU1, he gene conside ed he main esponsible o sulphi e ole ance by
egula ing he anspo o his molecule h ough he plasma ic memb ane
(A am and Bakalinsky, 1997), and in some s ains, also o hose genes ela ed
o sulphu me abolism.
Sec ion I – Resul s and Discussion
61
In any case, hose da a sugges ha each s ain o yeas has o con ain a ce ain
le el o SO2 wi hin he cell, and i can be pos ula ed ha such in e nal SO2 le el
will be co ela ed wi h he inal le el o SO2 emaining in he media a e
e men a ion. As one o he mos ob ious easons limi ing he e iciency o yeas
educ ases would be ha pa o he aldehydes we e p o ec ed by SO2, i can be
u he pos ula ed ha he a ios aldehyde/ alcohol and aldehyde/ acid should be
signi ican ly co ela ed o he inal le el o SO2 emaining in he media a e
e men a ion. This seems o be he case, as shown in Table 1 and pa icula ly in
Figu e 8, which highligh s h ee examples. In his Figu e, he a e age a ios o
he di e en samples (means o wo biological eplica es) a e seg ega ed by
yeas and ep esen ed e sus he inal con en in o al SO2 o he e men ed
media. Figu e 8a co esponds o phenylace aldehyde/ 2-phenyle hanol a io;
Figu e 8b o 3-me hylbu anal/ 3-me hylbu anoic acid a io and Figu e 8c o
me hional/ me hionol a io. I can be obse ed ha in he i s wo cases he h ee
s ains ollowed a simila dependency, so ha he ac ion o aldehyde emaining
0
5
10
15
20
25
30
35
40
45
50
L1 L2 L3
o al SO2, mg/L
To al SO2 o med by di e en yeas s ains a he end
o e men a ion
No SO₂ 30 mg/L SO₂
Figu e 7 – E ec s o yeas and o ex e nal SO2 on he inal le els o SO2: A e age inal SO2 le els ound
in samples e men ed wi h h ee ypes o s ain and wi h o wi hou addi ion o ex e nal SO2 (30 mg/L).
E o ba s a e s anda d e o s o he mean.

S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
62
pe uni o alcohol (Figu e 8a) o acid o med (Figu e 8b) is di ec ly p opo ional
o he le el o SO2, being he p opo ionali y cons an oughly independen o
he s ain o yeas . Howe e , in he hi d case, he p opo ionali y cons an
be ween he ac ion o aldehyde emaining pe uni o alcohol o med in yeas
3 is much smalle han hose o yeas s ains 1 and 2. This explains why he
o e all co ela ion coe icien be ween hese pa ame e s was no signi ican o
me hional (Table 1). As p e iously men ioned, his would be consis en wi h he
hypo hesis ha SO2 p esen wi hin he yeas cell (in bo h cy oplasm and
mi ochond ia) is he main ac o de e mining he inal le els o SAs a e
e men a ion. The s ong s ain dependency could, he e o e, be p ima ily due o
he yeas in insic me abolism o SO2.
Figu e 8 Aldehyde/alcohol o aldehyde/acid a ios and inal SO2 con en : Plo s showing he ela ionship
be ween impo an aldehyde/alcohol o aldehyde/acid a ios and he inal SO2 le els ound in samples
e men ed wi h h ee di e en yeas s ains. A) phenylace aldehyde/2-phenyle hanol a io; B) 3-
me hylbu anal/3-me hylbu anoic acid a io; C) me hional/ me hionol a io
0
0,5
1
1,5
2
2,5
010 20 30 40 50 60
Ra io
o al SO2
A. phenylace aldehyde/ phenyle hanol
0
10
20
30
40
50
60
010 20 30 40 50 60
Ra io
o al SO2
B. iso ale aldehyde/ iso ale ic acid
0
2
4
6
8
10
12
010 20 30 40 50 60
Ra io
o al SO2
C. me hional/ me hionol
L1 L2 L3
Sec ion I – Resul s and Discussion
63
3.3 Role o Zn on SA o ma ion
As p e iously men ioned, he e ec s o Zn on he o ma ion o SAs seems o be
seconda y (Table 1) only being signi ican in he cases o 3-me hylbu anal and
phenylace aldehyde in he expe imen in which he 0 le el o zinc was
conside ed. As shown in Table 3, he le els o hose aldehydes we e minimum
when le els o Zn a e 1 mg/L, becoming maxima a 10 and 0 mg/L. As in his
expe imen , le els o isobu anol, isoamyl alcohol and 2-phenyle hanol we e
signi ican ly smalle in condi ions o Zn s a a ion, some o he a ios aldehyde/
alcohol become a Zn s a a ion signi ican ly highe han hose obse ed a 1
mg/L. The same is obse ed in he aldehyde/ acid a io 3-me hylbu anal/ 3-
me hylbu anoic acid. Alcohol/ acid a ios we e also signi ican ly a ec ed by Zn
le els, signi ican ly inc easing wi h Zn le els (Table 3). These esul s sugges
Zn le els may ha e an e ec on he ADH ac i i ies o yeas which may ha e an
indi ec e ec on SAs o ma ion. In ac , Zn is an essen ial componen o many
dehyd ogenases (De Smid e al., 2008) which con ain a zinc-con aining ac i e
si e (Pe sson e al., 1993). Ou da a sugges ha he o e all ADH e iciency o
yeas s la gely dec eases in condi ions o comple e zinc s a a ion. These
obse a ions a e consis en wi h he known ac ha unde low zinc condi ion, a
egula o y me al- esponsi e p o ein al e s se e al S. ce e isiae me abolic
pa hways including ep essing some o he genes ha exp ess yeas ADH (Eide,
2009).
3.4 Fa y acids and hei e hyl es e s
As expec ed, le els o a y acids and o hei e hyl es e s we e signi ican ly
in luenced by he yeas s ain, as shown in Table 1. As seen in Table 2, yeas
s ain L1 p oduced maxima le els o he h ee acids and o wo o he e hyl
es e s, while L3 p oduced in mos cases he smalles le els. E ec s we e no
S ecke aldehydes a e no mal by-p oduc s o alcoholic e men a ion linked o yeas
sul i e me abolism
64
signi ican o e hyl hexanoa e, o which he es e i ica ion a io in L1 was
minimum.
Howe e , he mos ema kable e ec on le els o a y acids, hei e hyl es e s
and hei es e i ica ion a es a e played by Zn con en . As seen in Tables 1 and
3, Zn le els ha e a signi ican , bu no e y impo an e ec on he absolu e
le els o hexanoic and oc anoic acids, bu qui e in ense e ec s on he le els o
es e s and on he es e i ica ion a ios. As seen in Table 3, le els o e hyl
hexanoa e, oc anoa e and decanoa e eached maxima alues in condi ions o Zn
s a a ion o o low Zn le els in he i s expe imen . Maximum le els we e mo e
han 3-4 imes highe han he minimum le els. The e ec s on he es e i ica ion
a ios ollowed a simila end, wi h maxima le els in condi ions o Zn s a a ion
o o low Zn le els.
To he bes o ou knowledge, he ele an e ec s o Zn on e hyl es e s and
pa icula ly, on es e i ica ion a ios, ha e no been p e iously epo ed. By using
genomic and ansc ip omic analysis i has been known o a ime ha he
ac i i ies o all he enzymes o he cy idine diphospha e diacylglyce ol (CDP-
DAG) pa hway, he majo ou e in he syn hesis o phospholipids, a e dec eased
in Zn-limi ed cells (Iwanyshyn e al., 2004) while a di e en se o enzymes
ela ed o an al e na i e ou e o syn hesis known as Kennedy pa hway, displays
mo e ac i i y (Ke s ing and Ca man, 2006; So o and Ca man, 2008). I.e., i is
well es ablished ha in low Zn condi ions he e is a me abolic emodelling in he
syn hesis o phospholipids (Eide, 2009), and he e o e i should no be su p ising
ha he le els o a y acids and hei e hyl es e s, which a e by-p oduc s o such
syn hesis, change.
Sec ion I – Resul s and Discussion
65
10 mg/L 5 mg/L 1 mg/L 10 mg/L 1 mg/L 0 mg/L
o al SO2*20.3 ± 3.7b13.6 ± 3.5a13.8 ± 2.5 a ---
aldehydes
2-me hylp opanal - - - - - -
3-me hylbu anal*---33 ± 1.8b24.5 ± 1.6a32.8 ± 3.4b
2-me hylbu anal - - - - - -
me hional - - - - - -
phenylace aldehyde*---25.5 ± 5b17 ± 2a24.1 ± 4.9b
usel alcohols
isobu anol*---30 ± 8b26.6 ± 5.4b21.6 ± 4.8a
isoamyl alcohol - - - 207 ± 29b202 ± 16b163 ±17a
me hionol - - - - - -
2-phenyle hanol*17.7 ± 2a20.6 ± 1.4ab 23.1 ± 1.2b27.2 ± 2.7b27.9 ± 1.4b23.8 ± 1a
iso-acids
2-me hylp opanoic acid - - - - - -
3-me hylbu anoic acid 1.2 ± 0.1a1.5 ± 0.1b1.5 ± 0.2 b ---
aldehyde/ alcohol a io
2-me hylp opanal/isobu anol - - - - - -
3-me hylbu anal/isoamyl alcohol*---0.2 ± 0.0b0.1 ± 0.0a0.2 ± 0.0b
me hional/ me hionol - - - 5.1 ± 0.3a5.6 ± 0.5a6.7 ± 1b
phenylace aldehyde/2-phenyle hanol*---1 ± 0.3b0.6 ± 0.0a1 ± 0.2b
aldehyde/ acid a io
2-me hylp opanal/2-me hylp opanoic acid - - - - - -
3-me hylbu anal/3-me hylbu anoic acid*---26.9 ± 4.2b17.3 ± 2.8a23.6 ± 4.4b
isobu anol/2-me hylp opanoic acid - - - 20.4 ± 1.4b18.8 ± 1.9ab 13.6 ± 1.2a
isoamyl alcohol/3-me hylbu anoic acid - - - 151 ± 8b136 ± 13ab 108 ± 8a
2-me hylbu anal/3-me hylbu anal• - - - - - -
medium chain a y acids
hexanoic acid• - - - 1.5 ± 0.2b1.4 ± 0.2ab 1.3 ± 0.2a
oc anoic acid 2.7 ± 0.2b2.3 ± 0.2a2.9 ± 0.4b---
decanoic acid• - - - - - -
es e s
isoamyl ace a e - - - - - -
e hyl hexanoa e 0.1 ± 0.0a0.1 ± 0.0a0.4 ± 0.0b0.2 ± 0.0a0.2 ± 0.0a0.7 ± 0.1b
e hyl oc anoa e• 0.2 ± 0.0a0.1 ± 0.0a0.5 ± 0.1b0.3 ± 0.1a0.3 ± 0.1a1 ± 0.1b
e hyl decanoa e• 0.1 ± 0.0a0.1 ± 0.0a0.3 ± 0.1b0.1 ± 0.0a0.1 ± 0.0a0.4 ± 0.1b
a ios
e hyl hexanoa e/hexanoic acid 8.7 ± 0.8a9.2 ± 0.6a31.9 ± 4b12.7 ± 1.1a12.4 ± 1a47.5 ± 7.7b
e hyl oc anoa e/oc anoic acid 5.0 ± 0.7a4.8 ± 0.4a14.8 ± 0.7b8.2 ± 1.1a8.2 ± 1.2a29.3 ± 3.6b
e hyl decanoa e/decanoic acid 15.7 ± 1.5a11.7 ± 0.8a44.8 ± 5.2b11.4 ± 1.2a12.7 ± 0.6a42 ± 3.8b
No SO2
30ppm SO2
Table 3 A e age le els o o al SO2, a oma compounds and o some ele an a ios a ending o he Zn
le els o he mus in he wo expe imen s. Concen a ion da a a e in mg/L excep aldehydes which a e in
µ
g/L. Cases showing signi ican in e ac ions Zn x yeas a e ma ked wi h • o * o he expe imen s wi hou
o wi h SO2, espec i ely. Signi ican di e ences a ending o Duncan es a e indica ed wi h le e s a-c
being “a” he lowes a e age alue.

Sec ion II
Roles o yeas on he o ma ion and e olu ion o he a oma
o Riesling and Ga nacha wines
In oduc ion and Me hodology
Chap e 1 E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces
on he o ma ion and u he e olu ion o Riesling a oma
Chap e 2 The oles o yeas s on he o ma ion and e olu ion o Ga nacha wine
a oma
Chap e 3 Obse a ions, ques ions and conclusions de i ed om he compa ison
be ween a ie ies
In oduc ion and Me hodology
Sec ion II - In oduc ion
77
1. Sec ion II – In oduc ion
Fo long, he a oma o wine has been known o ha e di e en o igins and has
been adi ionally di ided in o g ape-de i ed compounds, e men a i e o aged
ela ed compounds. None heless, a oma o ma ion is an in e ac i e and complex
p ocess, which implies ha hese h ee g oups a e no necessa ily seg ega ed. In
ac , g ape composi ion deeply modula es he ac i i y o mic oo ganisms
ca ying ou e men a ion so ha e men a i e p o iles can be s ongly dependen
on he a ie y o g ape (Fe ei a e al., 1996; He nández-O e e al., 2002)).
Simila ly, he de elopmen and e olu ion o wine a oma wi h ime and hence,
i s aging po en ial is ela ed o a complex a ay o chemical p ocesses ac ing on
compounds de i ed om he g ape o o med du ing e men a ion (Fe ei a and
San Juan, 2012).
The majo i y o a oma compounds in neu al g apes a e p esen as non- ola ile
p ecu so s. Many o hem a e conjuga es o an a oma molecule and a non-
ola ile and wa e -soluble molecule, such as a glycoside. They ep esen a
undamen al sou ce o p ecu so s o a ie al a oma compounds in wine.
Glycosidic p ecu so s a e o med by one o mo e suga moie ies (glycones)
linked o an aglycone which can o igina e a ola ile odo an upon elease. To
da e, se e al aglycones ha e been iden i ied and a y om s aigh chain
alcohols o e penoids, shikimic acid me aboli es o no isop enoids. Glycosidic
p ecu so s a e linked o he o ma ion o impo an a ie al compounds such as
linalool, ge aniol, b-damascenone, a-ionone and b-ionone o 1,1,6- ime hyl-
1,2-dihyd onaph halene (TDN) (Hjelmeland and Ebele , 2015; Win e hal e and
Rouse , 2001; Win e hal e and Skou oumounis, 1997; Zoecklein e al., 1999).
S udies wi h glycosidic p ecu so s we e i s ca ied ou wi h a oma ic a ie ies
such as Musca , bu also wi h Riesling o i s high con en in se e al o he

Sec ion II - In oduc ion
78
compounds p e iously men ioned, and o he pa icula in e es a oused by he
de elopmen wi h ime o desc ip o s such as lo al, ci us o ke osene (Fische ,
2007; Simpson, 1978; Simpson and Mille , 1983; Win e hal e e al., 1990;
Zoecklein e al., 1999). On he o he hand, li le is known abou he ele ance o
glycosidic p ecu so s o o m he a oma o ed a ie ies, specially Ga nacha.
This a ie y has been desc ibed wi h black ui , chocola e and e en lowe y
no es howe e he compounds in ol ed in he o ma ion o hese desc ip o s a e
no ully known (Lopez e al., 2004).
In o de o o m an odo an , he aglycone has o be eleased om he glycoside
ei he by slow acid hyd olysis a wine pH o by enzyma ic hyd olysis. This las
can be ca ied ou by enzymes om he plan , bu i akes mainly place by he
ac ion o he di e en mic oo ganisms ca ying ou e men a ion. Alcohols and
mono e penes ha e been iden i ied as compounds ha could be di ec ly eleased
om glycosidic p ecu so s (Wa e house e al., 2016; Williams e al., 1980,
1993). None heless, some ele an a oma compounds a e no o med by s aigh
hyd olysis o he glycosidic bound be ween he suga and he aglycone bu a e
o med a e u he spon aneous chemical ea angemen s o he aglycone. This
is he case o some ele an a oma compounds de i ed om ca o enoids like
no isop enoids (Fische , 2007; Mendes-Pin o, 2009; Wa e house e al., 2016;
Win e hal e e al., 1990; Win e hal e and Rouse , 2001; Win e hal e and
Skou oumounis, 1997).
Di icul ies a ise since he hyd olysis o one pa icula p ecu so can o igina e
di e en compounds and one speci ic odo an can be o en o med om di e en
p ecu so s. This makes ha linking a oma compounds wi h speci ic p ecu so s
is a he di icul ask (Waldmann and Win e hal e , 1992; Win e hal e and
Skou oumounis, 1997; Zoecklein e al., 1999). Mo eo e , he e is also e idence
ha mos glycosides a e no hyd olysed du ing winemaking o ha hey a e
Sec ion II - In oduc ion
79
hyd olysed yielding non- ola ile compounds like polyols, which by u he
chemical ea angemen will yield he a oma molecule (Williams e al., 1980;
Zoecklein e al., 1999). This implies ha e men a ion can ha e a qui e complex
se o e ec s on he a oma po en ial o wine, and ha many o hese e ec s will
no be iden i ied bu a e long ime. Fu he mo e, hose e ec s can be u he
in luenced by s o age condi ions, oxygen con ac and p esence o lees
(Zoecklein e al., 1998, 1999), which adds mo e di icul ies in he a ionaliza ion
o he e ec s o e men a ion on a ie al a oma. The e iciency o he hyd olysis
due o enzyma ic ac i i y is highly s ain dependen , since he glycosidase
ac i i ies o di e en s ains can di e bo h in in ensi y and in he ange o ac i e
subs a es. Addi ional ac i i ies can be ound in non-Saccha omyces gene a,
which has leaded o he de elopmen o e men a ions combining cul u es o
non-Saccha omyces and Saccha omyces ce e isiae sequen ially inocula ed.
These s a egies ha e been shown o modula e a oma o ma ion and o ha e
po en ial o p oduce wines o highe quali y and complexi y, which has been
a ibu ed o he di e en abili ies o elease ola iles om g ape p ecu so s
(Beni o e al., 2015; Esc ibano e al., 2017; Padilla e al., 2016). Howe e , ew
s udies ha e ac ually been ca ied ou wi h ex ensi e analysis o he a oma
compounds o med, and ew less ha e aken in o conside a ion he e ec s o
aging, hence he e a e ye many open ques ions ega ding he ole o yeas in
wine a oma o ma ion and e olu ion, especially in wha conce ns hei ac ion on
g ape de i ed p ecu so s.
Sec ion II - In oduc ion
80
2. Goals
The p esen se o s udies in ends o b ing some ligh in o he oles played by
yeas s, in he o ma ion and de elopmen o a ie al and e men a i e a oma o
wine. Fo his, a speci ic esea ch in ol ing sequen ial e men a ions wi h
di e en yeas s ains, syn he ic mus con aining eal ac ions o p ecu so s
om wo g ape a ie ies and di e en aging imes has been ca ied ou . The aims
o he s udy a e:
1. To de e mine he hie a chy o ac o s (yeas , p ecu so s, ime) a ec ing
wine a oma p o ile.
2. To assess he speci ic e ec s linked o he p esence o a oma p ecu so s
on wine a oma and on i s e olu ion wi h ime.
3. To assess he e ec s o yeas on a ie al and e men a i e a omas and on
he e olu ion o a oma wi h ime.
4. To de i e gene al p ac ical conclusions abou he possibili ies o
modula e wine a oma using sequen ial e men a ions.
Sec ion II - Me hodology
81
3. Sec ion II – Me hodology
3.1 Reagen s and s anda ds
Dichlo ome hane (DCM), e hanol and me hanol (≥ 99%) Dis o-Pes icide esidue
g ade we e supplied by Me ck (Da ms ad , Ge many). Milli-Q® sys em om
Millipo e (Me ck, Ge many).
2-bu anol (≥ 99%), 4-me hyl-2-pen anol (99%), 4-hyd oxi-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 and 2-oc anol (99.5%), 3-
oc anone (99%) and 3,4-dime hylphenol (99%) we e used as in e nal s anda ds
o mino and ace compounds analysis and we e pu chased om Me ck.
The chemical s anda ds used in his s udy we e supplied by Me ck wi h pu i y ³
98%. TDN was syn hesised by Synchem UG & Co wi h a pu i y o 80%.
An alkane solu ion in dichlo ome hane (C7-C28) was used o calcula e
app oxima e linea e en ion index o analy es.
3.2 Glycosidic p ecu so s ex ac ion
3.2.1 G ape p ocessing
The glycosidic p ecu so ac ions we e ob ained du ing ha es 2016 and
app oxima ely 23 Kg o g apes we e ob ained o each a ie y. Riesling g apes
we e ob ained in Neus ad an de Weins asse, Ge many and Ga nacha g apes
we e gi en by Bodegas Román om D.O. Campo de Bo ja, Spain.
The g apes we e c ushed by ee and cold mace a ed o 24 hou s in he case o
Riesling and 48 hou s in he case o Ga nacha, in he p esence o La azymâCL
(La o , F ance). The di e ences o mace a ion ime ela e wi h he ac ha
whi e and ed winemaking ha e o iginally di e en mace a ion pe iods, and hus
his app oxima es eal winemaking o he small-scale expe imen s. To p o ec
Sec ion II - Me hodology
88
Table 4 Mass spec a ions selec ed o quan i y mino and ace compounds using GC-MS.
Compounds RT m/z
E hyl es e s and ace a es
E hyl isobu y a e 7.5 71a, 116
E hyl 2-me hylbu y a e 12.0 57a, 102
E hyl 3-me hylbu y a e 13.10 88a, 115, 70
E hyl 4-me hylpen anoa e 24.15 88a, 101
E hyl cyclohexanoa e 45.55 83a, 101, 156
Isobu yl ace a e 9.71 56a, 73
Phenyle hyl ace a e 79.90 91a
No isop enoids
Rose oxide 39.73/ 40.93 139a, 154
Vi ispi ane*52.8/ 53.08 192a, 93, 121, 171
Riesling ace al*59.9 138a, 125, 133
b-damascenone 79.86 69a, 190
a-ionone 72,4 121a, 93, 192
b-ionone 77.08 177a, 192
1,1,6-T ime hyl-1,2-dihyd onaph halene (TDN) 66.48 157a, 142, 172
Mono e penes
Linalool 55.01 71a, 93, 121
a- e pineol 64.05 93a, 121, 136
Ge aniol 72.63 69a, 123
b-ci onellol 68.13 69a, 81, 123
Lac ones
d-nonalac one 81.81 85a, 100
d-decalac one 87.36 85a, 100
Whiskylac one 74.56/ 78.18 99a, 114
Cinnama es
E hyl dihyd ocinnama e 74.54 178a, 133
E hyl cinnama e 86.80 131a, 176
Vola ile phenols
Guaiacol 73.5 109a, 124
o-c esol 81.16 108a, 79
m-c esol 85.35 108a, 79
4-e hylguaiacol 82.17 137a, 152
Eugenol 88.73 164a, 149
E-isoeugenol 96.83 164a, 149
4-e hylphenol 89.33 107a, 122
4-p opylguaiacol 85.96 137a, 166
4- inylguaiacol 90.14 150a, 135
4- inylphenol 99.04 120a, 91
2.6-dime hoxyphenol 93.27 154a, 139
4-allyl-2,6-dime hoxyphenol 104.87 194a, 119
Vanillin de i a es
Vanillin 105.85 151a, 152, 123
Ace o anillone 108.83 166a, 123
Sy ingaldehyde 127.15 182a, 181, 167
*Compounds en a i ely quan i ied using alkanes o de e mine he e en ion index;
a Quan i a i e agmen s m/z

Sec ion II - Me hodology
89
Fo Vi ispi ane and Riesling ace al a comme cial s anda d was no a ailable
hus, hei iden i ica ion was made using m/z and e en ion index om
bibliog aphy e e ences (Loscos e al., 2007) in SCAN mode as well as injec ion
o alkanes o calcula e e en ion index in a DB-wax column.
3.5 Da a ea men
Rela i e a eas we e ob ained by di iding he ion peak a ea o he analy e by he
a ea o he co esponding in e nal s anda d. Those a eas we e ans o med in o
concen a ions by in e pola ion in he calib a ion g aphs buil by he analysis o
calib a ed samples. Da a p ocessing was made using Mic oso Excel Visual
Basic o applica ion (VBA) simple coding.
Analysis o a iance (ANOVA) was made on he compounds wi h a ea abo e
he limi o quan i ica ion, assessing he ac o s p esence o p ecu so ac ion,
yeas s ain and accele a ed aging ime as well as he bina y in e ac ions
(p esence o p ecu so s x yeas s ain and yeas s ain x aging ime). P incipal
Componen Analysis and Sca e plo s we e used o analyse he da a. These
analyses we e pe o med using XLSTAT (Addinso , 2018 e sion).
The comple e da a se o each g ape a ie y was also analyzed by P incipal
Componen Analysis (PCA) o assess he hie a chy o ac o s a ec ing he
a oma o ma ion.
All g aphics we e made using Mic oso Excel, 2016 e sion.
The da a ob ained o each a ie y we e analysed indi idually in chap e s I and
II and a u he compa a i e analysis was pe o med in Chap e III.
Bibliog aphy
90
4. Bibliog aphy
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assimilable ni ogen de iciencies du ing alcoholic e men a ion in oenological
condi ions. Jou nal o Fe men a ion and Bioenginee ing 70, 246–252.
Beni o, S., Ho mann, T., Laie , M., Lochbühle , B., Schü le , A., Ebe , K., e
al. (2015). E ec on quali y and composi ion o Riesling wines e men ed by
sequen ial inocula ion wi h non-Saccha omyces and Saccha omyces ce e isiae.
Eu opean Food Resea ch and Technology 241, 707–717.
Esc ibano, R., González-A enzana, L., Ga ijo, P., Be lanas, C., López-Al a o,
I., López, R., e al. (2017). Sc eening o enzyma ic ac i i ies wi hin di e en
enological non-Saccha omyces yeas s. Jou nal o Food Science and Technology
54, 1555–1564.
Fe ei a, V., Fe nández, P., and Cacho, J. F. (1996). A S udy o ac o s a ec ing
wine ola ile composi ion and i s applica ion in disc iminan analysis. LWT -
Food Science and Technology 29, 251–259.
Fe ei a, V., and San Juan, F. (2012). “Fla o o Wine,” in Food Fla o s:
Chemical, Senso y, and Technological P ope ies.
Fische , U. (2007). “Wine A oma,” in Fla ou s and F ag ances: Chemis y,
Biop ocessing and Sus ainabili y, ed. R. G. Be ge (Be lin, Heidelbe g: Sp inge
Be lin Heidelbe g), 241–267.
He nández-O e, P., Cacho, J., and Fe ei a, V. (2002). Rela ionship be ween
a ie al amino acid p o ile o g apes and wine a oma ic composi ion.
expe imen s wi h model solu ions and chemome ic S udy. Jou nal o
ag icul u al and ood chemis y 50, 2891–2899.
Hjelmeland, A. K., and Ebele , S. E. (2015). Glycosidically Bound Vola ile
A oma Compounds in G apes and Wine: A Re iew. Ame ican Jou nal o
Enology and Vi icul u e 66.
Lopez, R., Azna , M., Cacho, J., and Fe ei a, V. (2002). De e mina ion o mino
and ace ola ile compounds in wine by solid-phase ex ac ion and gas
ch oma og aphy wi h mass spec ome ic de ec ion. Jou nal o Ch oma og aphy
A 966, 167–177.
Lopez, R., Ezpele a, E., Sánchez, I., Cacho, J., and Fe ei a, V. (2004). Analysis
o he a oma in ensi ies o ola ile compounds eleased om mild acid
hyd olysa es o odou less p ecu so s ex ac ed om Temp anillo and G enache
Bibliog aphy
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g apes using gas ch oma og aphy-ol ac ome y. Food Chemis y 88, 95–103.
Loscos, N., He nandez-O e, P., Cacho, J., and Fe ei a, V. (2007). Release and
o ma ion o a ie al a oma compounds du ing alcoholic e men a ion om
non lo al g ape odo less la o p ecu so s ac ions. Jou nal o Ag icul u al and
Food Chemis y 55, 6674–6684.
Mendes-Pin o, M. M. (2009). Ca o enoid b eakdown p oduc s he—
no isop enoids—in wine a oma. A chi es o Biochemis y and Biophysics 483,
236–245.
OIV Compendium o In e na ional Me hods o Analysis o Wines and Mus s (2
ol.). oi .in . A ailable a : h p://www.oi .in /en/ echnical-s anda ds-and-
documen s/me hods-o -analysis/compendium-o -in e na ional-me hods-o -
analysis-o -wines-and-mus s-2- ol.
O ega, C., Lopez, R., Cacho, J., and Fe ei a, V. (2001). Fas analysis o
impo an wine ola ile compounds De elopmen and alida ion o a new
me hod based on gas ch oma og aphic – lame ionisa ion de ec ion analysis o
dichlo ome hane mic oex ac s. 923, 205–214.
Padilla, B., Gil, J. V., and Manzana es, P. (2016). Pas and u u e o non-
Saccha omyces yeas s: om spoilage mic oo ganisms o bio echnological ools
o imp o ing wine a oma complexi y. F on ie s in Mic obiology 7, 1–20.
Simpson, R. F. (1978). A oma and composi ional changes in wine wi h
oxida ion, s o age and ageing. Vi is, 274–287.
Simpson, R. F., and Mille , G. C. (1983). A oma composi ion o aged Riesling
wine. Vi is 22, 51–63.
Waldmann, D., and Win e hal e , P. (1992). Iden i ica ion o a no el is ispi ane
p ecu so in Riesling wine. Vi is 31, 169–174.
Wa e house, A. L., Sacks, G. L., and Je e y, D. W. (2016). Unde s anding wine
chemis y. Wiley.
Williams, P. J., Se on, M. A., and Ma inos, V. A. (1993). Hyd oly ic la o
elease om non- ola ile p ecu so s in ui s, wines and some o he plan -
de i ed oods. in, 283–290.
Williams, P. J., S auss, C. R., and Wilson, B. (1980). Hyd oxyla ed linalool
de i a i es as p ecu so s o ola ile mono e penes o musca g apes. Jou nal o
Ag icul u al and Food Chemis y 28, 766–771.
Win e hal e , P., and Rouse , R. (2001). “Ca o enoid-De i ed A oma
Compounds: An In oduc ion,” in Ca o enoid-De i ed A oma Compounds, eds.
P. Win e hal e and R. L. Rouse (Washing on, DC: Ame ican Chemical
Bibliog aphy
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Socie y), 1–17.
Win e hal e , P., Se on, M. A., and Williams, P. J. (1990). Vola ile C13 -
no isop enoid compounds, in Riesling wine a e gene a ed om mul iple
p ecu so s. Ame ican Jou nal o Enology and Vi icul u e 41, 277–283.
Win e hal e , P., and Skou oumounis, G. K. (1997). “Glycoconjuga ed a oma
compounds: occu ence, ole and bio echnological ans o ma ion,” in Ad ances
in Biochemical Enginee ing/ Bio echnology (Be lin, Heidelbe g: Sp inge Be lin
Heidelbe g), 73–105.
Zoecklein, B. W., Hackney, C. H., Duncan, S. E., and Ma cy, J. E. (1999). E ec
o e men a ion, aging and he mal s o age on o al glycosides, phenol- ee
glycosides and ola ile compounds o Whi e Riesling (Vi is ini e a L.) wines.
Jou nal o Indus ial Mic obiology and Bio echnology 22, 100–107.
Zoecklein, B. W., Jasinski, Y., and McMahon, H. (1998). E ec o e men a ion,
aging, and aging su lie on o al and phenol- ee Riesling (Vi is ini e a L.)
glycosides. Jou nal o Food Composi ion and Analysis 11, 240–248.

Chap e 1
E ec s o sequen ial inocula ion wi h di e en non-
Saccha omyces on he o ma ion and u he e olu ion o
Riesling wine a oma
Sec ion II – Chap e 1
97
Chap e 1 - E ec s o sequen ial e men a ion wi h
di e en non-Saccha omyces on he o ma ion and
e olu ion o Riesling a oma
1. Resul s and discussion
The expe imen al app oach ollowed in his wo k makes i possible o iden i y
he a oma compounds ha a e exclusi ely o med om componen s p esen in
he g ape glycosidic p ecu so ac ion, di e encing hem om hose which a e
o med exclusi ely due o yeas me abolism. The o me will be only ound in
samples con aining p ecu so s, e men ed o non- e men ed -acid-hyd olysis
con ols. The la e will be ound in all he e men ed samples, ega dless o he
p esence o glycosidic p ecu so ac ion.
The app oach also allows he iden i ica ion o di e en ial e ec s o yeas s on
he o ma ion o a ie al compounds, i s by compa ing e men ed samples wi h
he un e men ed con ols con aining jus p ecu so s om which, a oma
compounds a e o med by acid hyd olysis and second by compa ing he samples
e men ed wi h di e en yeas s.
Finally, he app oach adds a ime a iable, since wines ha e been submi ed o
accele a ed aging allowing he iden i ica ion o di e en aging pa e ns linked o
he p esence o yeas .
The wines we e i s ly cha ac e ized acco ding o hei classical oenological
pa ame e s and hen a compa a i e analysis o he ola ile composi ion among
he di e en con ols and wines spiked wi h glycosidic p ecu so s was made.
E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
104
samples made only wi h S. ce e isiae a e ound in he middle o he plane and
no clea ly sepa a ed om hose made wi h T. delb ueckii, which a e a hei le
side. Finally, all samples e men ed wi h L. The mo ole ans ha e nega i e sco es
in he i s componen . The in luence o aging ime is also seen in he igu e,
wi h aged samples ha ing highe sco es in he second componen . Howe e , he
p esence o p ecu so s has no clea e ec in he ep esen a ion, meaning ha he
con en s in e men a i e ola iles a e no highly a ec ed by he p esence o
p ecu so s in he e men ing mus .
The a iable loading plo , gi en in he uppe pa o he igu e, shows ha nea ly
all componen s ha e posi i e loadings in he i s componen , which is
pa icula ly co ela ed wi h ola ile a y acids, hei e hyl es e s and wi h he
ace a es o usel alcohols. Only 1-bu anol, e hyl lac a e and 1-hexanol, keep a
nega i e co ela ion wi h he i s componen .
This, ce ainly, indica es ha samples e men ed wi h P. kluy e i ha e he
highes le els o mos ola ile compounds, no ably o a y acids, hei e hyl
es e s and o he ace a es o highe alcohols.
As o he second componen , i is posi i ely co ela ed wi h he e hyl es e s o
b anched acids, wi h g-bu y olac one and b-phenyle hanol and nega i ely
co ela ed wi h he ace a es o usel alcohols and wi h bu y ic acid.
Figu es 11 o 13 include a selec ion o plo s showing he e olu ion wi h ime o
he di e en compounds in he wines e men ed wi h di e en yeas s, in o de
o acili a e he in e p e a ion o esul s.
Figu e 11, gi es he plo s wi h he e olu ion o ace a es and o he es e s and
acids. As can be seen, isoamyl ace a e and phenyle hyl ace a e a e ound a much
highe le els in samples e men ed wi h P. kluy e i. I is also ob ious ha le els
o ace a es a e sligh ly, bu signi ican ly, highe in samples no con aining

Sec ion II – Chap e 1
105
p ecu so s, sugges ing ha he ace yl ans e ase ac i i y has been nega i ely
in luenced by he ac ion o p ecu so s. Ne e heless, ecen ly e men ed
samples con ain mo e han 2.5 mg/L o phenyle hyl ace a e, an amoun
exceeding, by a , he odou h eshold o his compound. Le els o isoamyl
ace a e a e, howe e , no pa icula ly la ge. Bo h compounds ollow a dec easing
end wi h ime, since he acid-alcohol/es e equilib ium is displaced owa ds he
dissocia ed o m. Ye , le els o phenyle hyl ace a e a e 5 weeks o aging a e
high enough o ha e high senso y implica ions.
Hexanoic and decanoic acids, as well as hei co esponding e hyl es e s a e also
illus a ed in Figu e 11. Also, in his case, i is e iden ha samples e men ed
by P. kluy e i ha e he highes le els, al hough di e ences a e no as ma ked as
o ace a es. None heless, le els ollowed he o de P. kluy e i < Saccha omyces
< T. delb ueckii < L. The mo ole ans. Le els o he e hyl es e s a e no
pa icula ly high, bu his can be pa ly a ibu ed o he la ge e apo a ion a e o
hese compounds when e men a ion is ca ied ou in small olumes.
Ne e heless, le els o he co esponding a y acids a e no mal-high, p omo ing
he e hyl es e s con en o emain cons an wi h ime, con a ily o he case o
ace a es.
E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
106
The mos ele an highe alcohols in wine can be seen in Figu e 12. In all cases,
yeas s ains had a signi ican e ec , bu he ou come is compound dependen .
In he case o isoamyl alcohol, wines e men ed wi h S. ce e isiae con ained
signi ican ly highe le els, while wines e men ed wi h P. kluy e i had he
0
100
200
300
400
500
600
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Isoamyl ace a e
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
500
1000
1500
2000
2500
3000
3500
4000
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Phenyleh yl ace a e
S. ce e isiae P. kluy e i T. delb ueckii L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
50
100
150
200
250
300
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
E hyl hexanoa e
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
500
1000
1500
2000
2500
3000
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Hexanoic acid
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
50
100
150
200
250
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
E hyl decanoa e
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
500
1000
1500
2000
2500
0 1 2 5
Concen a ion (mg/L)
Measu emen ime (week)
Decanoic acid
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
Figu e 11 Yeas and aging e ec s on ace a e es e s, a y acids and hey e hyl es e s – e olu ions wi h ime
o wo ace a es, wo e hyl es e s and hei co esponding a y acids (µg/L) acco ding o he p esence (PR)
o absence (CTL) o p ecu so s and o he yeas gene a ha ca ied ou e men a ion; samples we e aken
a e e men a ion – 0 and 1, 2 and 5 weeks o accele a ed aging. Ace a es and e hyl es e s show dec easing
endencies while acids show inc easing endencies wi h ime. P. kluy e i ou s ands in he p oduc ion o
es e s and o hexanoic acid.
Sec ion II – Chap e 1
107
smalles le els. On he con a y, o me hionol and b-phenyle hanol he le els
ollow he o de P. kluy e i > T. delb ueckii > S. ce e isiae > L. The mo ole ans;
In he case o isobu anol, he pa e n o all yeas is simila o me hionol and b-
phenyle hanol wi h he excep ion o S. ce e isiae which can p oduce equi alen
le els o hose o P. kluy e i. Le els o hese compounds emain ai ly s able
du ing aging, as can be seen in he igu e.
Rega ding e hyl es e s o b anched acids, hese compounds a e o med by slow
es e i ica ion o hei co esponding acids. Acco dingly, le els o he e hyl es e s
a e close o 0 in he ecen ly e men ed samples, inc easing wi h ime, as shown
in Figu e 13. Le els o isobu y ic acid a e ai ly s able wi h ime. Rega ding
yeas ap i ude o o m hese wo compounds, he o de was T. delb ueckii > P.
kluy e i > S. ce e isiae > L. The mo ole ans and, as obse ed in he case o
ace a es, samples e men ed wi hou p ecu so s ha e highe con en s han hose
spiked wi h Riesling p ecu so s. Fe men a i e compounds a e by-p oduc s o
yeas seconda y me abolisms. Se e al a oma compounds a e o med in ou es
ela ed o yeas amino acids me abolism and a second g oup o yeas lipid
me abolism. A hi d g oup, he ace a es, a e ela ed wi h bo h me abolic ou es
since hei p oduc ion is o med by ace yl-CoA ans e ases ac ing on highe
alcohols which a e p oduced du ing he syn hesis o amino acids. Thus, i is
e iden ha hese ou es a e highly s ain-dependen and a e also in luenced by
he in e ac ion be ween non-Saccha omyces and S. ce e isiae (Flee , 2003). In
ac , and as shown in Figu e 9 and 10, despi e ha all wines we e sequen ially
inocula ed wi h S. ce e isiae, non-Saccha omyces in oduced a majo sou ce o
a oma a iabili y on all main me abolic ou comes.
E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
108
0
100
200
300
400
500
600
700
800
0 1 2 5
Concen a ion (µg/L)
Measu emen (week)
Isobu y ic acid
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
20
40
60
80
100
120
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
E hyl isobu y a e
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
Figu e 12 Yeas and aging e ec s on b anched acids and e hyl es e s - one o he b anched acids and i s
co esponding e hyl es e e olu ions wi h ime (µg/L) acco ding o he p esence (PR) o absence (CTL) o
p ecu so s and o he yeas gene a ha ca ied ou e men a ion; samples we e aken a e e men a ion
and a e 1, 2 and 5 weeks o accele a ed aging. Each yeas s ains shows simila o ma ion pa e ns o
bo h compounds. The es e inc eases con inuously and he acid is s able wi h ime.
0
10000
20000
30000
40000
50000
60000
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Isoamyl alcohol
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
1000
2000
3000
4000
5000
6000
7000
0 1 2 5
Concen a ion (µg/L)
Measu emen (week)
Me hionol
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
10000
0 1 2 5
Concen a ion (µg/L)
Measu emen (week)
β-phenyle hanol
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
0
2000
4000
6000
8000
10000
12000
0 1 2 5
Concen a ion (µg/L)
Measu emen (week)
Isobu anol
PR S. ce e isiae PR P. kluy e i PR T. delb ueckii PR L. he mo ole ans
CTL S. ce e isiae CTL P. kluy e i CTL T. delb ueckii CTL L. he mo ole ans
Figu e 13 Yeas and aging e ec s on usel alcohols - majo usel alcohols e olu ion wi h ime (µg/L)
acco ding o he p esence (PR) o absence (CTL) o p ecu so s and o he yeas gene a ha ca ied ou
e men a ion; samples we e aken a e e men a ion and a e 1, 2 and 5 weeks o accele a ed aging. L.
he mo ole ans p oduces lowes le els o hese compounds. Fusel alcohols a e qui e s able du ing ime.
Sec ion II – Chap e 1
109
The e ec s o P. kluy e i on he le els o a y acids, hei e hyl es e s and
pa icula ly on he ace a es o highe alcohols, clea ly show ha hese me abolic
ou es ha e been much p omo ed in he p esence o his yeas , al hough S.
ce e isiae has ca ied ou mos pa o he e men a ion (Padilla e al., 2016). On
he con a y, he p esence o L. he mo ole ans has he opposi e e ec , s ongly
limi ing he numbe o es e s, ace a es and a y acids p oduced du ing
e men a ion. On he o he hand, L. he mo ole ans has an ou s anding capaci y
o p oduce e hyl lac a e as can be seen in he a iables plo o Figu e 9. Tha is
mos likely linked o his yeas epo ed ap i ude o o m lac ic acid du ing
alcoholic e men a ion (Beni o e al., 2015; Gobbi e al., 2013; Kapsopoulou e
al., 2007). Samples e men ed wi h T. delb ueckii on i s side, show e men a i e
ola ile p o iles close o hose o S. ce e isiae. Rema kably, he h ee non-
Saccha omyces esul ed in wines wi h smalle le els o isoamyl alcohol
compa ed wi h e men a ions ca ied en i ely ou by S. ce e isiae. This educ ion
may ha e senso y ele ance since his compound is a s ong supp esso o wine
ui y and woody no es (de-la-Fuen e-Blanco e al., 2016). Fu he mo e, hese
da a sugges ha sequen ial inocula ion o non-Saccha omyces yeas s ains
leads o wines wi h highe a oma complexi y (Esc ibano e al., 2017; Jolly e al.,
2014). Indeed, all samples whe e non-Saccha omyces yeas we e inocula ed had
inal a oma con en ai ly di e en om S. ce e isiae e ealing ha his
me hodology has epe cussions on he inal wine p o ile. The ac ha wines
e men ed wi h P. kluy e i ha e o e all highe es e con en and abo e hei
odou h eshold is likely o esul in mo e ui y and lowe y-like wines. On he
o he hand, L. he mo ole ans s ongly limi s he le els o e men a i e odo an s
o med, sugges ing ha i can be an impo an modula o o wine ac ile
p ope ies, due o i s abili y o p oduce lac ic acid and i s de i a i es (Beni o e
al., 2015; Swiege s e al., 2005) and also ha is able o p oduce wines in which
non- e men a i e no es will be mo e easily pe cei ed.

E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
110
1.4 Va ie al compounds in Riesling
Figu e 14 summa izes he PCA ca ied ou on a ie al a oma compounds ound
in he da a se . Only samples con aining p ecu so s, e men ed o no , we e
included in he analysis. The wo i s componen s e ain nea ly a 60% o he
o iginal a iance. As can be seen in he plo , he ac o mos in luen ial in he
loca ion o samples in he plane is aging ime, ollowed by e men a ion and he
yeas s ain. The eshly e men ed samples ha e he mos nega i e sco es in he
i s componen , wi h a single excep ion o S. ce e isiae.
On he con a y, samples wi h 5 weeks o accele a ed aging ha e highes sco es
in he i s componen . Addi ionally, un e men ed samples ha e he highes
sco es o he second componen con a ily o samples e men ed wi h S.
ce e isiae wi h equi alen aging imes, which ha e he smalles sco es on he
same axis. The plo sugges s ha in his case, samples e men ed wi h S.
ce e isiae a e he mos di e en o he pu e a ie al a oma ob ained by simple
acid hyd olysis, while he in e en ion o non-Saccha omyces yeas s c ea es
a ie al wine p o iles mo e simila o hose obse ed by simple acid hyd olysis.
A look a he sample loading plo o Figu e 6, e eals ha aging is ela ed o
ge aniol and linalool dec eases and wi h TDN and inylphenols inc eases.
Mo eo e , he exis ence o e men a ion leads o inc eased le els o
ace o anillone and minima o linalool, a- e pineol and b-damascenone.
Figu es 15-17 ep esen he e olu ion wi h ime o a ie al compounds in he
wines e men ed wi h he di e en yeas s ains h oughou ime. Looking a he
di e en plo s, i should be no ed ha in some ele an cases, no ably hose o
linalool, a- e pineol, Riesling ace al, b-damascenone and ge aniol, un e men ed
con ols con ained always highe le els han e men ed samples (Figu es 15 and
16).
Sec ion II – Chap e 1
111
TDN
β-damascenone
Linalool
α- e pineol
β-ci onellol
Ge aniol
Guaiacol
4- inylguaiacol
2-6-
dime hoxyphenol
E-isoeugenol
4- inylphenol
anillin
ace o anillone
β-ionone
E hyl cinnama e
-1
-0,75
-0,5
-0,25
0
0,25
0,5
0,75
1
-1 -0,75 -0,5 -0,25 00,25 0,5 0,75 1
F2 (25,43 %)
F1 (34,31 %)
Va iables (axes F1 and F2: 59,74 %)
0
1
2
5
0
1
25
0
1
2
5
0
12
5
0
1
2
5
-5
-4
-3
-2
-1
0
1
2
3
4
5
-4 -3 -2 -1 01234567
F2 (25,43 %)
F1 (34,31 %)
Obse a ions (axes F1 and F2: 59,74 %)
AH L. he mo ole ans P. kluy e i S. ce e isiae T. delb ueckii
Figu e 14 P incipal Componen Analysis on a ie al compounds quan i ied in e men ed samples wi h
Riesling p ecu so ac ion, as well as un e men ed con ols spiked wi h p ecu so s. The plo shows he
p ojec ion o a iables ( op plo ) o samples (bo om) showing he wo biological eplica es- in he plane
o med by he i s wo componen s, which e ained 59.74 % o he o iginal a iance. Numbe s in he
samples e e o he weeks o anoxic s o age a 50°C.
E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
112
This pa e n was no eally expec ed and sugges s ha a la ge po ion o hese
compounds’ p ecu so s we e no eally glycosides, bu di e en polyols, which
by simple ea angemen in acid media, yielded he a oma compounds wi hin he
ew weeks be ween he p epa a ion o he syn he ic mus s and he ime o
analysis a e e men a ion. Al hough his migh a ec equally he e men ed and
un e men ed samples, since he ac ion is he same, he ola ile compounds
al eady p esen du ing e men a ion will ine i ably be pa ially co-e apo a ed
wi h CO2 p oduced du ing e men a ion, which helps explaining why le els in
e men ed samples a e consis en ly smalle .
Figu e 15 E olu ion wi h ime o he le els o he main mono e penes, linalool, ge aniol and α- e pineol,
in e men ed samples and un e men ed con ols con aining p ecu so s ex ac ed om Riesling g apes.
Samples we e aken a he end o e men a ion (0) and a e 1, 2 and 5 weeks o accele a ed aging. Wines
e men ed exclusi ely wi h S. ce e isiae we e used as con ols and e men a ions wi h non-Saccha omyces
s ains o P. kluy e i. T. delb ueckii and L. he mo ole ans we e sequen ially inocula ed wi h S. ce e isiae.
AH (acid hyd olysis) was used as un e men ed con ol o syn he ic wine spiked wi h he glycosidic p ecu so
ac ion. Da a o ge aniol con en in wines e men ed by P. Kluy e i a e gi en by a single sample.
0
20
40
60
80
100
120
140
160
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Linalool
AH S. ce e isiae P. kluy e i T. delb ueckii L. he mo ole ans
0
20
40
60
80
100
120
140
160
180
200
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
"
- e pineol
AH S. ce e isiae P. kluy e i T. delb ueckii L. he mo ole ans
0
10
20
30
40
50
60
0 1 2 5
Concen a ion (
!
g/L)
Measu emen ime (week)
Ge aniol
AH S. ce e isiae P. kluy e i T. delb ueckii L. he mo ole ans
Sec ion II – Chap e 1
113
The di e en cases will be b ie ly analysed and discussed.
Linalool and ge aniol a e qui e uns able compounds a wine pH and ha e a
gene al endency o dec ease du ing aging (Figu e 15). The dec easing a e is
mi iga ed since new molecules eleased om p ecu so s eplace he decomposed
ones. As a o emen ioned, he lowe le els in e men ed con ols could be
a ibu ed o he pa ial e apo a ion o ea ly o med a oma compounds du ing
e men a ion.
Di e ences be ween yeas s ains we e only mode a ely signi ican (see
supplemen a y da a) showing ha wines e men ed wi h P. kluy e i seem o ha e
he lowes con en o linalool, a- e pineol and ge aniol. The ac ion o such s ain
is no limi ed o a low e iciency in he hyd olysis o he p ecu so s, bu o he
ac ha p ecu so s we e p obably ans o med in o di e en compounds.
O he wise, a slowe a e o dec ease should ha e been obse ed. Da a also show
ha wines om L. he mo ole ans ha e signi ican ly highe le els o he h ee
a oma compounds a e 1 week o aging. This sugges s ha ei he he enzymes
exc e ed by his s ain du ing e men a ion o upon cell au olysis we e s ill ac i e
du ing accele a ed aging, o ha enzymes om his s ain we e pa icula ly
e icien a a oiding ans o ming p ecu so s in o molecules di e en o he
a ge ed odo an s.
Mos su p isingly, le els o ge aniol we e ound o inc ease in P. kluy e i wines
a e 2 weeks o aging. Le els in hese wines we e abo e 50 µg/L, he highes o
his compound obse ed. This peculia esul needs u he expe imen al
checking, since he analy ical esul s o one biological eplica e o ge aniol was
los .
The end ollowed by a- e pineol is di e en , since he compound inc eases o
a maximum le el be o e s a ing o dec ease, which is consis en wi h he ac
E ec s o sequen ial e men a ion wi h di e en non-Saccha omyces on he
o ma ion and u he e olu ion o Riesling a oma
120
Saccha omyces s ains ha e simila le els be ween hem and o hose ound in
he un e men ed con ol, may indica e low o e en absen hyd oxycinnama e
deca boxylase ac i i y in hese yeas s ains, sugges ing ha a la ge ac ion is
p oduced by simple acid hyd olysis o he glycoside.
None heless, no mally and especially in ed wines, inyl phenols ha e
dec easing endencies wi h ime due o hei eac i i y wi h an hocyanins. The
o ma ion o adduc s wi h e hanol has also been desc ibed as po en ial cause o
ola ile phenols dec eases in wine (Kennison e al., 2008; Wa e house e al.,
2016).

Sec ion II – Chap e 1
121
2. Conclusions
The sequen ial inocula ion e men a ion app oach used in his s udy in oduces
a la ge a iabili y in he pa e n o e men a i e compounds and in he wine
a oma p o ile. Wines ob ained by e men a ion wi h P. kluy e i we e he mos
dis inc , con aining highes le els o usel alcohol ace a es, o a y acids and o
hei e hyl es e s. On he o he hand, samples e men ed wi h L. he mo ole ans
con ained minima le els o hose compounds. Rema kably, all wines ob ained
by sequen ial inocula ion e men a ion con ained smalle le els o isoamyl
alcohol, which is a s ong odou supp esso .
Le els o some ele an a ie al a oma compounds we e ound a highe le els
in un e men ed con ols, sugges ing he exis ence o a pool o easily
hyd olysable p ecu so s, such as polyols, among Riesling p ecu so s. Among
e men ed samples, hose made wi h L. he mo ole ans con ained he highes
le els o mono e penes while hose e men ed wi h P. kluy e i con ined minima
le els o mono e penes and no isop enoids. The o ma ion o TDN is ime-
dependen bu i is also s ongly enhanced by e men a ion, sugges ing ha
hyd oly ic ac i i ies o yeas s a e essen ial o p oduce acid-hyd olysable
p ecu so s o his molecule. In compa ison wi h S. ce e isiae, wines made wi h
P. kluy e i ha e a lowe abili y o accumula e TDN. Mos ema kably, samples
e men ed wi h he h ee non-Saccha omyces yeas s had much smalle le els o
inylphenols.
In summa y, esul s p esen ed he e u he suppo ha a sequen ial inocula ion
e men a ion app oach can be success ully used no only o modula e wine
a oma bu o con ol i s e olu ion wi h ime.
Bibliog aphy
122
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Ca au, F. M., Medina, K., Boido, E., Fa ina, L., Gagge o, C., Dellacassa, E., e
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Daniel, M. A., Capone, D. L., Se on, M. A., and Elsey, G. M. (2009). Riesling
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Be lin Heidelbe g), 241–267.
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64, 9401–9411.
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Chap e 2
E ec s o sequen ial inocula ion wi h di e en non-
Saccha omyces on he o ma ion and u he
e olu ion o Ga nacha wine a oma
Sec ion II - Chap e 2
129
Chap e 2 - The oles o yeas s on he o ma ion and
e olu ion o Ga nacha wine a oma
1. Resul s and discussion
Resul s p esen ed in his chap e e e o e men a ions ca ied ou wi h syn he ic
mus con aining o no glycosidic p ecu so s ex ac ed om Ga nacha g apes.
The global goal o he chap e is o assess he oles o yeas s on he o ma ion
and e olu ion wi h ime o a oma compounds in Spanish Ga nacha wines.
Speci ic goals a e o be e de ine wha is a ie al a oma o Ga nacha wine and
o assess he in luence o yeas on i s de elopmen du ing wine aging.
A sequen ial inocula ion p o ocol was ollowed, meaning ha he s e ile
syn he ic mus s we e inocula ed i s wi h a non-Saccha omyces s ain and, a e
4 days, wi h S. ce e isiae o comple e he e men a ion, excep o one ial
exclusi ely e men ed wi h S. ce e isiae, which was kep as con ol. Hal o he
samples con ained only syn he ic mus wi h all he necessa y nu ien s and
elemen s o no mal yeas me abolism and he second hal we e addi ionally
spiked wi h he ac ion o glycosidic p ecu so s. Besides, un e men ed con ol
samples o syn he ic wine spiked wi h p ecu so s ac ion we e included in o de
o assess he ole o acid hyd olysis and, pa icula ly, enable a quan i a i e
compa ison be ween he e iciencies o acid e sus enzyma ic hyd olysis on he
a ie al a oma o ma ion. A ime a iable, in which wine was aged in a comple e
anoxic en i onmen o up o i e weeks a 50ºC was also included. This se -up
aims o u he unde s and he o igin and a e o he ola ile compounds
quan i ied in he wines, he ole o di e en yeas gene a and o slow hyd oly ic
p ocesses, igh a e e men a ion and du ing aging ime.
Impac o ineya d e sus cella mic obio a om di e en ha es s on he dis inc ion
o di e en Riesling ineya ds
232
glycosidic p ecu so s) no only in he o ma ion o a ie al cha ac e , bu also
and mos impo an ly in he cons uc ion o speci ic e oi nuances in wine.
Figu e 48 Va ie al a oma compounds in wines e men ed asep ically and in he cella om g apes om
5 di e en ineya ds in 2016 ha es – Wines e men ed om g apes handpicked asep ically om 7
di e en P alz egions and e men ed in asep ic condi ion ( ineya ds 1-7) o in cella (wg 1-7). Resul s a e
exp essed in
µ
g/L.
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
20
30
40
50
60
70
80
90
100
110
120
Linalool
Box plo s (Linalool)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
2
4
6
8
10
12
14
16
Ge aniol
Box plo s (Ge aniol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
2
4
6
8
10
12
Guaiacol
Box plo s (Guaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
100
200
300
400
500
600
700
4-Vinylguaiacol
Box plo s (4-Vinylguaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
200
400
600
800
1000
1200
4-Vinylphenol
Box plo s (4-Vinylphenol)

Sec ion III – Chap e 5
233
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
500
1000
1500
2000
2500
3000
Isoamyl ace a e
Box plo s (Isoamyl ace a e)
ineya d 1 ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
5
10
15
20
25
30
35
40
45
c-3-Hexenol
Box plo s (c-3-Hexenol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
100
200
300
400
500
600
700
800
900
Benzylic alcohol
Box plo s (Benzylic alcohol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
400
600
800
1000
1200
1400
1600
1800
Bu y ic acid
Box plo s (Bu y ic acid)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
100
200
300
400
500
600
700
800
900
1000
Isobu y ic acid
Box plo s (Isobu y ic acid)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
5
10
15
20
25
30
35
40
Iso ale aldehyde
Box plo s (Iso ale aldehyde)
Figu e 49 Fe men a i e a oma compounds in wines e men ed asep ically and in he cella om g apes
om 5 di e en ineya ds in 2016 ha es – Wines e men ed om g apes handpicked asep ically om 5
di e en P alz egions and e men ed in asep ic condi ion ( ineya ds 1-7) o in cella (wg 1-7). Resul s a e
exp essed in
µ
g/L.
Impac o ineya d e sus cella mic obio a om di e en ha es s on he dis inc ion
o di e en Riesling ineya ds
234
Figu e 50 Va ie al compounds in wines e men ed asep ically and in he cella om g apes om 5
di e en ineya ds in 2015 ha es – Wines e men ed om g apes handpicked asep ically om 5 di e en
P alz egions and e men ed in asep ic condi ion ( ineya d 1-5) o in cella (wg 1-5). Resul s a e exp essed
in
µ
g/L.
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
wg1 wg2 wg3 wg4 wg5
0
20
40
60
80
100
120
Linalool
Box plo s (Linalool)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
wg1 wg2 wg3 wg4 wg5
2
4
6
8
10
12
14
16
18
20
Ge aniol
Box plo s (Ge aniol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
wg1 wg2 wg3 wg4 wg5
0
2
4
6
8
10
12
14
16
18
Guaiacol
Box plo s (Guaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
wg1 wg2 wg3 wg4 wg5
0
100
200
300
400
500
600
700
4-Vinylguaiacol
Box plo s (4-Vinylguaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
wg1 wg2 wg3 wg4 wg5
0
200
400
600
800
1000
1200
4-Vinylphenol
Box plo s (4-Vinylphenol)
Sec ion III – Chap e 5
235
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
20
30
40
50
60
70
80
90
100
110
120
Linalool
Box plo s (Linalool)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
2
4
6
8
10
12
14
16
Ge aniol
Box plo s (Ge aniol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
2
4
6
8
10
12
Guaiacol
Box plo s (Guaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
100
200
300
400
500
600
700
4-Vinylguaiacol
Box plo s (4-Vinylguaiacol)
ineya d 1
ineya d 2
ineya d 3
ineya d 4
ineya d 5
ineya d 6
ineya d 7
wg1 wg2 wg4 wg5
wg6-202
wg6-77
wg6-775
wg7-209
wg7-30
0
200
400
600
800
1000
1200
4-Vinylphenol
Box plo s (4-Vinylphenol)
Figu e 51 Va ie al compounds in wines e men ed asep ically and in he cella om g apes om 5
di e en ineya ds in 2016 ha es – Wines e men ed om g apes handpicked asep ically om 5 di e en
P alz egions and e men ed in asep ic condi ion ( ineya d 1-5) o in cella (wg 1-5). Resul s a e exp essed
in
µ
g/L.
Impac o ineya d e sus cella mic obio a om di e en ha es s on he dis inc ion
o di e en Riesling ineya ds
236
2. Conclusions
Wines made om he same g ape a ie y and ineya d in wo consecu i e
ha es s show a comple ely di e en a oma composi ion. These di e ences a e
he likely esul o speci ic clima e condi ions o each yea , which closely ela e
o he ine managemen , including he deg ee o ma u i y o he g apes achie ed
in each yea and also, he li le changes in he winemaking p ac ices in oduce
as a esponse o he speci ic condi ions a which he g apes a i e o he cella
each ha es . These di e ences should ansla e in o bo h changes in ineya d
mic o lo a and changes in g ape be y composi ion, bo h o which can be
po en ially impo an a oma modula o s.
Al hough in his speci ic wo k, speci ically ela ed o he p elimina y s udy o
ola ile compounds, i is no possible o make a de ini i e assessmen o whe he
he obse ed changes a e due o chemical di e ences o o di e ences in he
mic obio a, he s udy shows a ele an in luence o he e oi wi hin each
in age.
Fe men a i e compounds such as es e s highly e lec he ha es e ec , whe eas
a ie al compounds o med om g ape glycosidic p ecu so s show simila
o ma ion pa e ns in wines om 2015 and 216 ha es . On he con a y,
poly unc ional me cap ans seem o be highly a ec ed and hei o ma ion
changes signi ican ly om ha es o ha es .
G apes picked om he same ineya ds and e men ed unde asep ic condi ions
o in comme cial cella en i onmen o igina e di e en wines in which cella
mic o lo a should ha e a majo e ec .

Bibliog aphy
238
3. Bibliog aphy
Capone, D. L., and Je e y, D. W. (2011). E ec s o anspo ing and p ocessing
Sau ignon blanc g apes on 3-me cap ohexan-1-ol p ecu so concen a ions.
Jou nal o Ag icul u al and Food Chemis y 59, 4659–4667.
Pa ke , M., Capone, D. L., F ancis, I. L., and He de ich, M. J. (2017). A oma
P ecu so s in G apes and Wine: Fla o Release du ing Wine P oduc ion and
Consump ion. Jou nal o Ag icul u al and Food Chemis y.
Pey o des Gachons, C., Tominaga, T., and Dubou dieu, D. (2000). Measu ing
he a oma ic po en ial o Vi is ini e a L. C . Sau ignon Blanc g apes by
assaying S -cys eine conjuga es, p ecu so s o he ola ile hiols esponsible o
hei a ie al a oma. Jou nal o Ag icul u al and Food Chemis y 48, 3387–
3391.
Pey o des Gachons, C., Tominaga, T., and Dubou dieu, D. (2002). Localiza ion
o S-cys eine conjuga es in he be y: e ec o skin con ac on a oma ic po en ial
o Vi is ini e a L. c . Sau ignon blanc mus . Ame ican Jou nal o Enology and
Vi icul u e 53, 144–146.
Ribé eau-Gayon, P., Glo ies, Y., Maujean, A., and Dubou dieu, D. (2006).
Handbook o Enology - Chemis y o wine, s abiliza ion and ea men s. 2nd ed.
Tominaga, T., Bal enweck-Guyo , R., Des Gachons, C. P., and Dubou dieu, D.
(2000). Con ibu ion o ola ile hiols o he a omas o whi e wines made om
se e al Vi is ini e a g ape a ie ies. Am. J. Enol. Vi ic. 51, 178–181.
Tominaga, T., Pey o des Gachons, C., and Dubou dieu, D. (1998). A New Type
o Fla o P ecu so s in Vi is ini e a L. c . Sau ignon Blanc: S -Cys eine
Conjuga es. Jou nal o Ag icul u al and Food Chemis y 46, 5215–5219.
Supplemen a y da a
Supplemen a y da a
248
3. Supplemen a y da a om Sec ion II: Chap e 1
S.d. Table 4 3-way ANOVA assessing he e ec o he ac o s: p esence o absence o p ecu so s, yeas
s ain, aging and hei in e ac ion on he ola ile composi ion o Riesling syn he ic wine. F and signi icance
a e indica ed o each ac o . Signi icance is exp essed as *: <0.0001-0.001***; 0.001-0.01**; 0.01-0.05*.
E hyl ace a e 18.0 *** 12.6 *** 1.3 n.s. 3.6 * 0.7 n.s.
Isoamyl ace a e 6.8 * 401.1 *** 24.2 *** 8.5 *** 21.6 ***
E hyl hexanoa e 0.0 n.s. 96.7 *** 1.7 n.s. 6.8 *** 0.5 n.s.
E hyl oc anoa e 0.0 n.s. 81.9 *** 1.8 n.s. 4.8 ** 1.2 n.s.
E hyl decanoa e 2.4 n.s. 28.8 *** 2.2 n.s. 2.6 * 1.1 n.s.
Isobu anol 0.1 n.s. 128.7 *** 0.5 n.s. 2.1 n.s. 0.4 n.s.
Isoamyl alcohol 0.7 n.s. 136.2 *** 0.1 n.s. 1.4 n.s. 0.1 n.s.
Me ionol 12.6 *** 221.4 *** 5.4 ** 1.5 n.s. 2.7 n.s.
β-Phenyle hanol 0.9 n.s. 111.8 *** 3.7 * 1.0 n.s. 1.5 n.s.
E hyl lac a e 0.5 n.s. 16.1 *** 5.6 ** 1.0 n.s. 1.0 n.s.
γ-Bu y olac one 1.7 n.s. 34.9 *** 45.2 *** 0.4 n.s. 2.6 n.s.
Bu y ic acid 0.6 n.s. 2.1 n.s. 1.1 n.s. 0.8 n.s. 0.7 n.s.
Isobu y ic acid 28.5 *** 141.5 *** 1.2 n.s. 6.2 *** 1.7 n.s.
Hexanoic acid 0.6 n.s. 367.6 *** 0.7 n.s. 12.5 *** 0.6 n.s.
Oc anoic acid 0.8 n.s. 340.0 *** 1.1 n.s. 9.6 *** 0.9 n.s.
Decanoic acid 1.0 n.s. 18.9 *** 1.1 n.s. 0.1 n.s. 0.9 n.s.
E hyl isobu y a e 4.4 * 48.4 *** 76.7 *** 2.1 n.s. 9.8 ***
Isobu yl ace a e 19.6 *** 916.5 *** 57.6 *** 23.4 *** 61.2 ***
E hyl 2-me hylbu y a e 0.2 n.s. 43.4 *** 91.1 *** 1.8 n.s. 10.2 ***
Phenyle hyl ace a e 27.9 *** 1197.0 *** 61.7 *** 33.2 *** 68.9 ***
γ-nonalac one 10.9 ** 145.0 *** 3.6 * 2.1 n.s. 0.8 n.s.
γ-decalac one 1.1 n.s. 328.5 *** 7.6 *** 2.1 n.s. 1.9 n.s.
TDN 34.2 *** 1.2 n.s. 11.1 *** 1.2 n.s. 0.4 n.s.
β-damascenone 1100.1 *** 24.4 *** 8.2 *** 34.6 *** 0.8 n.s.
Linalool 78.0 *** 4.8 ** 15.3 *** 5.0 ** 0.5 n.s.
α- e pineol 186.8 *** 4.4 ** 12.0 *** 4.4 ** 0.4 n.s.
β-ci onellol 26.4 *** 3.8 ** 23.8 *** 1.0 n.s. 1.6 n.s.
Ge aniol 46.6 *** 3.7 * 1.4 n.s. 4.1 ** 2.1 n.s.
4- inylguaiacol 122.7 *** 1.8 n.s. 14.3 *** 1.9 n.s. 0.7 n.s.
4- inylphenol 166.9 *** 8.3 *** 6.4 *** 8.3 *** 2.7 **
anillin 52.4 *** 7.3 *** 6.3 *** 8.0 *** 2.3 *
ace o anillone 1102.0 *** 50.3 *** 1.5 n.s. 47.3 *** 1.5 n.s.
Aging
Yeas
P ecu so s
P ecu so s*yeas
Yeas *Aging

Supplemen a y da a
249
Mos o CTL CTL CTL CTL
Le adu a
Aging 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5
E hyl ace a e 0 0 0 0 0 0 0 0 5625 ± 103 9936 ± 1171 13483 ± 1107 16193 ± 1708 30264 ± 262 29553 ± 1016 13523 ± 13523 23211 ± 778 28147 ± 996 29245 ± 131 27402 ± 1792 20699 ± 950 23011 ± 22656 22606 ± 21452 32724 ± 2401 24590 ± 592
Isoamyl ace a e 0 0 0 0 0 0 0 0 30.6 ± 2.9 19.0 ± 3.8 14.8 ± 0.7 14.9 ± 4.5 32.4 ± 1.2 24.1 ± 0.0 17.1 ± 5.9 15.3 ± 1.0 492 ± 35 385 ± 38 416 ± 69 120 ± 19 395 ± 27 295 ± 65 283 ± 1 95.7 ± 13.9
Isobu yl ace a e 1.75 1.50 1.18 1.26 2.39 1.60 1.79 1.39 17.2 ± 1.2 16.3 ± 1.1 16.9 ± 1.5 18.1 ± 1.2 20.5 ± 1.6 20.9 ± 0.3 20.7 ± 0.1 21.1 ± 1.0 263 ± 1 211 ± 3 165 ± 6 59.0 ± 5.0 194 ± 4 148 ± 12 128 ± 6 51.5 ± 2.4
Phenyle hyl ace a e 0.00 0.75 0.00 0.00 4.84 4.54 4.92 3.75 22.7 ± 1.1 17.1 ± 0.6 13.1 ± 0.6 9.4 ± 0.5 23.6 ± 0.6 18.7 ± 0.2 15.0 ± 0.2 11.4 ± 0.1 3727 ± 55 3078 ± 53 2477 ± 119 864 ± 55 2713 ± 62 2112 ± 215 1780 ± 16 671 ± 12
E hyl hexanoa e 0 0 0 0 0 0 0 0 70.1 ± 4.3 71.4 ± 31.0 65.1 ± 1.5 66.2 ± 12.0 94.7 ± 11.9 96.7 ± 10.1 86.8 ± 14.7 86.3 ± 3.9 123 ± 5 167 ± 14 129 ± 20 131 ± 3 162 ± 25 180 ± 62 171 ± 30 143 ± 28
E hyl oc anoa e 0 0 0 0 0 0 0 0 104 ± 5 73.1 ± 30.8 99.3 ± 28.9 0.0 ± 0.0 138 ± 20 103 ± 4 111 ± 8 118 ± 10 168 ± 13 160 ± 17 189 ± 30 150 ± 1 146 ± 28 144 ± 13 148 ± 10 172 ± 33
E hyl decanoa e 0 0 0 0 0 0 0 0 34.2 ± 1.2 66.1 ± 11.4 54.3 ± 5.8 46.2 ± 27.5 47.9 ± 6.4 62.3 ± 13.6 48.0 ± 9.7 51.3 ± 3.4 87.9 ± 6.4 144 ± 47 92.1 ± 10.1 52.8 ± 4.7 141 ± 25 130 ± 83 73.8 ± 1.1 72.1 ± 17.7
E hyl isobu y a e 0 0 0 0 0 0 0 0 1.1 ± 1.1 10.0 ± 0.5 21.9 ± 1.9 38.4 ± 6.1 1.4 ± 0.0 6.2 ± 0.4 13.6 ± 0.6 25.1 ± 2.7 4.2 ± 0.7 26.9 ± 1.6 48.8 ± 2.3 57.1 ± 1.2 2.7 ± 0.0 16.5 ± 0.8 33.3 ± 0.7 51.2 ± 17.0
E hyl 2-me hylbu y a e 0 0 0 0 0 0 0 0 0.0 ± 0.0 1.0 ± 0.1 2.3 ± 0.0 3.6 ± 0.5 0.6 ± 0.1 0.8 ± 0.0 2.0 ± 0.0 3.0 ± 0.5 0.0 ± 0.0 1.2 ± 0.1 2.4 ± 0.1 5.5 ± 0.1 0.6 ± 0.1 1.0 ± 0.1 2.5 ± 1.0 4.4 ± 0.9
E hyl iso ale a e 0 0 0 0 0 0 0 0 0.0 ± 0.0 0.9 ± 0.1 1.8 ± 0.2 3.4 ± 0.3 0.1 ± 0.0 0.7 ± 0.0 1.4 ± 0.1 2.8 ± 0.1 0.1 ± 0.0 0.9 ± 0.0 1.4 ± 0.1 3.9 ± 0.2 0.1 ± 0.0 0.6 ± 0.0 1.3 ± 0.1 3.1 ± 0.1
E hyl lac a e 0 0 0 0 0 0 0 0 100 ± 9 437 ± 9 668 ± 27 867 ± 68 110 ± 3 474 ± 18 841 ± 29 1158 ± 14 105 ± 24 413 ± 110 618 ± 136 1032 ± 311 117 ± 11 424 ± 19 785 ± 146 1226 ± 105
Isobu anol 0 0 0 0 0 0 0 0 8115 ± 946 7823 ± 1038 8226 ± 1900 7602 ± 966 8645 ± 441 8753 ± 201 9917 ± 197 9198 ± 159 8251 ± 510 9662 ± 714 8694 ± 186 7136 ± 552 9407 ± 3 9009 ± 1814 8744 ± 372 8409 ± 30
1-Bu anol 0 0 0 0 0 0 0 0 60.0 ± 0.5 55.5 ± 4.0 55.2 ± 3.8 61.4 ± 2.4 63.8 ± 0.1 57.3 ± 6.6 58.4 ± 7.8 57.4 ± 8.8 47.1 ± 1.6 58.8 ± 13.3 63.6 ± 8.5 55.7 ± 0.7 54.0 ± 2.4 52.2 ± 0.7 53.8 ± 0.9 73.0 ± 18.4
Isoamyl alcohol 0 0 0 0 0 0 0 0 42922 ± 5794 41979 ± 6773 43341 ± 8181 41487 ± 5854 44279 ± 2897 44837 ± 1801 47300 ± 1647 45515 ± 2382 31823 ± 1978 33373 ± 3472 31011 ± 2881 31604 ± 2823 34994 ± 266 33848 ± 3496 35941 ± 1470 36035 ± 874
1-Hexanol 0 0 0 0 0 0 0 0 11.7 ± 0.1 7.5 ± 7.5 12.3 ± 0.1 9.9 ± 9.9 128 ± 4 137 ± 1 132 ± 3 141 ± 1 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 5.4 ± 5.4 27.8 ± 0.4 31.3 ± 2.0 35.6 ± 0.5 39.1 ± 0.7
Me ionol 0 0 0 0 0 0 0 0 2960 ± 112 3319 ± 360 3110 ± 439 3192 ± 220 3390 ± 166 3555 ± 64 3604 ± 38 3630 ± 107 3490 ± 75 4190 ± 251 4436 ± 217 4855 ± 23 3940 ± 29 4165 ± 860 5577 ± 733 5002 ± 90
β-Phenyle hanol 0 0 0 0 0 0 0 0 4913 ± 521 5379 ± 717 5172 ± 788 5336 ± 643 5206 ± 139 5633 ± 257 5533 ± 40 5601 ± 215 5157 ± 53 5652 ± 391 6235 ± 352 7316 ± 42 5364 ± 188 5663 ± 580 7835 ± 983 7353 ± 46
γ-Bu y olac one 0 0 0 0 0 0 0 0 215 ± 41 932 ± 148 959 ± 98 965 ± 248 192 ± 8 870 ± 68 1025 ± 108 1050 ± 58 128 ± 86 875 ± 16 1021 ± 157 979 ± 50 240 ± 22 850 ± 99 1288 ± 233 1169 ± 124
γ-nonalac one 0.65 0 0 0.47 0 0 0 0 2.6 ± 1.6 4.7 ± 0.1 4.6 ± 0.0 4.9 ± 0.3 4.9 ± 0.1 4.9 ± 0.2 5.0 ± 0.1 5.3 ± 0.2 4.4 ± 0.1 5.0 ± 0.0 5.2 ± 0.1 5.2 ± 0.1 5.0 ± 0.3 5.0 ± 0.1 5.4 ± 0.4 5.5 ± 0.5
γ-decalac one 0 0 0 0 0 0 0 0 2.8 ± 0.4 2.9 ± 0.2 3.1 ± 0.1 3.0 ± 0.1 2.3 ± 0.1 2.8 ± 0.0 2.7 ± 0.2 2.9 ± 0.1 3.3 ± 0.3 3.9 ± 0.1 3.7 ± 0.2 4.2 ± 0.2 3.2 ± 0.1 3.7 ± 0.2 3.9 ± 0.3 4.3 ± 0.3
Bu y ic acid 0 0 0 0 0 0 0 0 285 ± 39 161 ± 27 104 ± 30 133 ± 42 360 ± 150 185 ± 15 210 ± 74 248 ± 3 2395 ± 2151 217 ± 6 442 ± 73 230 ± 82 278 ± 114 348 ± 186 262 ± 38 239 ± 12
Isobu y ic acid 0 0 0 0 0 0 0 0 198 ± 16 219 ± 19 184 ± 25 233 ± 20 111 ± 2 98.7 ± 20.7 133 ± 5 141 ± 13 493 ± 24 500 ± 6 638 ± 54 332 ± 111 309 ± 9 272 ± 7 339 ± 1 294 ± 20
Hexanoic acid 0 0 0 0 0 0 0 0 1037 ± 113 1049 ± 63 1055 ± 15 1060 ± 78 1203 ± 61 1193 ± 18 1245 ± 58 1255 ± 43 2030 ± 48 2017 ± 179 2100 ± 146 2069 ± 9 2208 ± 134 2169 ± 222 2609 ± 37 2380 ± 51
Oc anoic acid 0 0 0 0 0 0 0 0 2321 ± 122 2223 ± 200 2077 ± 97 2574 ± 69 2614 ± 95 2728 ± 33 2605 ± 155 2936 ± 113 3664 ± 374 3559 ± 229 3948 ± 460 4032 ± 134 3960 ± 214 3754 ± 358 4185 ± 171 4602 ± 213
Decanoic acid 0 0 0 0 0 0 0 0 960 ± 126 606 ± 63 610 ± 18 1067 ± 234 852 ± 149 741 ± 114 893 ± 105 1033 ± 17 1035 ± 131 796 ± 45 1271 ± 308 1417 ± 18 982 ± 58 1177 ± 414 1327 ± 5 1736 ± 226
TDN 0.84 0.27 0.27 0.20 1.61 5.06 10.4 37 1.5 ± 0.2 0.2 ± 0.0 0.3 ± 0.0 0.5 ± 0.5 1.4 ± 0.0 14.3 ± 0.7 47.4 ± 0.3 96.6 ± 0.4 0.5 ± 0.0 0.7 ± 0.0 0.0 ± 0.0 0.4 ± 0.0 0.5 ± 0.0 5.3 ± 0.8 20.1 ± 2.9 46.9 ± 6.4
β-damascenone 0 0 0 0 6.28 6.72 7.37 5.92 0.0 ± 0.0 0.1 ± 0.0 0.3 ± 0.0 0.3 ± 0.1 2.5 ± 0.0 3.4 ± 0.2 3.8 ± 0.0 4.5 ± 0.1 0.4 ± 0.0 0.5 ± 0.0 0.4 ± 0.0 0.3 ± 0.0 1.6 ± 0.0 2.6 ± 0.2 2.4 ± 0.2 3.0 ± 0.3
β-ionone 0.28 0.56 0.57 0.08 0.31 0.33 0.71 0.10 0.5 ± 0.0 0.2 ± 0.0 0.6 ± 0.1 0.5 ± 0.0 0.4 ± 0.0 0.2 ± 0.0 0.5 ± 0.0 0.4 ± 0.0 0.8 ± 0.0 0.3 ± 0.0 0.4 ± 0.0 0.1 ± 0.0 0.8 ± 0.1 0.3 ± 0.0 0.6 ± 0.2 0.3 ± 0.4
E hyl cinnama e 00.13 0.00 0.02 0.05 0.09 0.10 0.15 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.1 ± 0.0 0.1 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
Linalool 0.76 2.10 1.98 0.29 150 114 76.7 9.37 4.2 ± 0.3 1.2 ± 0.1 2.8 ± 0.1 2.1 ± 0.1 62.9 ± 0.7 42.8 ± 1.3 7.1 ± 0.1 2.5 ± 0.0 2.2 ± 0.1 1.4 ± 0.1 1.9 ± 0.0 0.5 ± 0.0 46.3 ± 0.7 31.5 ± 4.9 9.4 ± 1.7 1.9 ± 1.4
α- e pineol 0.65 0.92 1.07 0.46 78.5 148 178 158 1.1 ± 0.1 1.5 ± 0.0 0.9 ± 0.5 1.0 ± 0.2 12.5 ± 12.0 139 ± 4 111 ± 1 50.1 ± 1.2 1.0 ± 0.0 0.9 ± 0.0 0.9 ± 0.1 0.3 ± 0.0 20.2 ± 0.4 73.1 ± 1.1 73.1 ± 2.7 26.1 ± 0.7
β-ci onellol 0.23 0.71 0.63 00.43 0.60 0.86 02.3 ± 0.2 0.5 ± 0.0 0.8 ± 0.1 0.8 ± 0.2 6.9 ± 0.1 2.1 ± 0.0 1.1 ± 0.1 0.7 ± 0.1 1.3 ± 0.0 0.5 ± 0.0 0.6 ± 0.0 0.0 ± 0.0 4.5 ± 0.3 1.6 ± 0.1 1.1 ± 0.2 0.3 ± 0.5
Ge aniol 01.75 0 0 38.7 36.2 26.6 1.88 2.6 ± 1.4 0.0 ± 0.0 0.4 ± 0.0 0.3 ± 0.3 9.6 ± 0.1 12.1 ± 0.0 1.7 ± 0.0 0.3 ± 0.3 0.4 ± 0.1 0.0 ± 0.0 0.5 ± 0.0 0.0 ± 0.0 5.5 ± 0.0 8.9 ± 1.6 51.2 38.5 ± 9.5
Guaiacol 0.06 0.04 0.06 0.00 0.41 0.38 0.58 1.86 0.3 ± 0.0 0.1 ± 0.0 0.1 ± 0.0 0.3 ± 0.1 2.2 ± 0.1 2.1 ± 0.1 0.9 ± 0.3 5.3 ± 3.8 0.2 ± 0.0 0.1 ± 0.0 0.1 ± 0.0 0.4 ± 0.1 0.9 ± 0.1 0.7 ± 0.1 1.0 ± 0.5 2.4 ± 0.6
4- inylguaiacol 27.3 44.5 28.6 55.5 348 746 1404 5335 16.0 ± 9.9 27.1 ± 7.3 11.3 ± 4.4 67.0 ± 44.9 1902 ± 283 2016 ± 38 2207 ± 100 3496 ± 618 61.4 ± 2.1 24.0 ± 0.3 34.4 ± 8.1 106 ± 28 656 ± 11 682 ± 7 1506 ± 382 2705 ± 16
2-6-dime hoxyphenol 0.16 0 0 0.09 0 0 0 0 0.6 ± 0.2 0.1 ± 0.1 0.0 ± 0.0 0.5 ± 0.5 1.5 ± 1.5 0.0 ± 0.0 0.0 ± 0.0 4.1 ± 0.7 0.2 ± 0.2 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.3 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 1.4 ± 0.1
E-isoeugenol 0 0 0 0 0 0.19 0 0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.4 ± 0.0 0.0 ± 0.0 0.2 ± 0.2 0.5 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.1 ± 0.1
4- inylphenol 14.6 15.9 14 36.1 364 944 1318 4299 149 ± 23 26.0 ± 1.5 16.7 ± 3.6 54.0 ± 36.8 4599 ± 237 2255 ± 78 2079 ± 214 2791 ± 523 70.7 ± 0.5 18.4 ± 1.3 35.2 ± 13.6 124 ± 26 1165 ± 85 825 ± 9 1065 ± 51 1774 ± 153
anillin 1.58 0.67 00.78 13.16 14.3 41.9 91 1.8 ± 0.7 2.5 ± 1.2 1.0 ± 0.0 3.7 ± 2.5 9.8 ± 1.6 9.5 ± 0.0 10.0 ± 0.8 23.5 ± 9.5 1.6 ± 0.2 0.9 ± 0.1 1.1 ± 0.1 1.8 ± 0.4 7.2 ± 0.4 7.4 ± 1.4 6.6 ± 0.3 8.7 ± 1.4
ace o anillone 0.76 0 0 0 5.02 5.75 8.30 13.9 0.0 ± 0.0 1.0 ± 0.7 0.0 ± 0.0 2.3 ± 1.8 58.4 ± 0.8 60.7 ± 0.1 58.8 ± 4.7 58.2 ± 2.2 1.5 ± 1.5 0.3 ± 0.3 0.2 ± 0.2 0.0 ± 0.0 32.9 ± 0.9 29.4 ± 1.4 30.8 ± 1.4 32.1 ± 1.7
Con ol
PR
AH
PR
S. ce e isiae
P. kluy e i
CTL
PR
S.d. Table 5 A e age concen a ion o ola iles measu ed abo e he limi o quan i ica ion in con ol wines and hose spiked wi h glycosidic p ecu so s om
Riesling g apes in un e men ed con ols (Acid Hyd olysis) and in wines e men ed wi h S. ce e isiae. P. kluy e i. T. delb ueckii and L. he mo ole ans.
Supplemen a y da a
251
Mos o
Le adu a
Aging 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5
E hyl ace a e 15005 ± 370 14768 ± 1645 13958 ± 244 6059 ± 5986 43321 ± 1201 38412 ± 772 28832 ± 84 17938 ± 1208 13398 ± 2392 12579 ± 2117 11574 ± 1377 10963 ± 479 26460 ± 12517 33153 ± 3367 26992 ± 3771 17095 ± 3153
Isoamyl ace a e 17.2 ± 2.2 16.9 ± 1.5 11.1 ± 1.4 0.0 ± 0.0 10.8 ± 1.9 11.0 ± 3.5 11.5 ± 1.9 0.0 ± 0.0 16.9 ± 2.4 12.3 ± 1.1 13.0 ± 3.3 0.0 ± 0.0 11.0 ± 1.4 16.3 ± 3.6 13.8 ± 2.0 12.5 ± 0.1
Isobu yl ace a e 22.4 ± 0.2 21.2 ± 1.0 18.1 ± 0.9 15.6 ± 0.6 20.2 ± 5.4 18.0 ± 4.8 10.0 ± 1.6 13.8 ± 3.3 9.2 ± 0.4 8.0 ± 0.8 7.5 ± 0.3 7.5 ± 0.3 5.9 ± 1.8 6.4 ± 1.3 11.9 ± 5.5 7.5 ± 1.3
Phenyle hyl ace a e 48.9 ± 3.1 39.4 ± 3.0 32.7 ± 2.1 20.6 ± 1.1 48.0 ± 5.2 36.1 ± 4.0 16.6 ± 10.7 17.3 ± 1.2 3.6 ± 0.0 2.9 ± 0.1 2.6 ± 0.0 2.5 ± 0.1 5.3 ± 1.3 4.4 ± 0.9 18.7 ± 14.4 5.0 ± 0.5
E hyl hexanoa e 76.3 ± 6.1 67.1 ± 7.9 80.2 ± 7.6 59.6 ± 1.9 37.8 ± 21.0 39.7 ± 11.5 34.7 ± 5.9 0.0 ± 0.0 39.9 ± 1.5 37.9 ± 5.2 31.5 ± 2.4 32.3 ± 9.8 26.0 ± 5.6 43.7 ± 17.8 35.1 ± 6.0 21.5 ± 1.1
E hyl oc anoa e 77.5 ± 10.3 66.7 ± 12.1 26.5 ± 26.5 0.0 ± 0.0 0.0 ± 0.0 27.5 ± 27.5 49.7 ± 19.8 0.0 ± 0.0 37.3 ± 6.1 27.0 ± 27.0 40.9 ± 4.7 34.7 ± 5.4 0.0 ± 0.0 20.9 ± 0.7 26.5 ± 9.4 25.1 ± 3.3
E hyl decanoa e 62.1 ± 0.2 53.8 ± 11.0 60.4 ± 1.5 74.8 ± 15.2 0.0 ± 0.0 48.0 ± 1.1 26.6 ± 1.7 0.0 ± 0.0 19.0 ± 19.0 43.2 ± 10.4 0.0 ± 0.0 27.5 ± 1.1 0.0 ± 0.0 0.0 ± 0.0 16.5 ± 1.1 0.0 ± 0.0
E hyl isobu y a e 4.0 ± 0.3 25.7 ± 3.3 44.2 ± 2.0 93.2 ± 4.3 4.9 ± 1.2 27.0 ± 6.3 25.8 ± 9.8 77.9 ± 9.3 2.2 ± 0.4 5.2 ± 5.2 7.9 ± 7.9 31.2 ± 1.6 1.1 ± 1.1 6.4 ± 1.5 28.2 ± 17.9 26.6 ± 5.0
E hyl 2-me hylbu y a e 0.6 ± 0.0 1.7 ± 0.0 2.7 ± 0.2 5.9 ± 0.5 0.6 ± 0.0 1.8 ± 0.2 1.4 ± 0.5 5.2 ± 0.7 0.3 ± 0.3 0.3 ± 0.3 1.0 ± 0.2 1.1 ± 0.1 0.5 ± 0.0 0.4 ± 0.0 1.5 ± 0.8 2.3 ± 0.4
E hyl iso ale a e 0.0 ± 0.0 0.6 ± 0.0 1.1 ± 0.1 2.4 ± 0.2 0.0 ± 0.0 0.6 ± 0.1 0.6 ± 0.1 2.1 ± 0.5 0.0 ± 0.0 0.1 ± 0.1 0.2 ± 0.2 0.6 ± 0.1 0.0 ± 0.0 0.1 ± 0.1 0.7 ± 0.4 0.4 ± 0.4
E hyl lac a e 93.9 ± 10.8 335 ± 5 602 ± 49 1067 ± 184 119 ± 34 363 ± 93 588 ± 169 944 ± 176 364 ± 179 2313 ± 1097 3475 ± 1413 5141 ± 2459 1852 ± 1664 1060 ± 156 1839 ± 399 2910 ± 606
Isobu anol 7668 ± 225 7857 ± 293 8389 ± 482 8074 ± 654 7332 ± 1567 7865 ± 1237 7408 ± 1272 7344 ± 1344 5254 ± 632 5136 ± 244 5104 ± 197 5394 ± 277 4451 ± 452 4663 ± 962 4894 ± 917 4227 ± 675
1-Bu anol 93.2 ± 1.8 105 ± 10 113 ± 2 110 ± 0 104 ± 6 102 ± 4 92.1 ± 13.5 93.5 ± 14.9 110 ± 2 115 ± 3 102 ± 16 96.9 ± 10.8 109 ± 1 110 ± 0 106 ± 4 91.3 ± 10.1
Isoamyl alcohol 35329 ± 1164 36262 ± 1071 37246 ± 957 39467 ± 1418 33266 ± 5029 34338 ± 4879 33867 ± 4951 34361 ± 4040 33012 ± 1798 33355 ± 102 32447 ± 744 32648 ± 730 34520 ± 5166 33537 ± 4542 35077 ± 5092 34306 ± 5380
1-Hexanol 12.1 ± 0.3 13.7 ± 0.8 14.0 ± 0.8 19.4 ± 2.0 130 ± 4 129 ± 1 137 ± 5 138 ± 3 12.0 ± 0.4 17.6 ± 2.1 14.4 ± 2.5 16.1 ± 0.5 159 ± 10 156 ± 2 156 ± 3 164 ± 5
Me ionol 4114 ± 1 3871 ± 346 4315 ± 146 5400 ± 930 4355 ± 71 4220 ± 38 4446 ± 79 5587 ± 952 1483 ± 54 1616 ± 90 1307 ± 18 1616 ± 12 2404 ± 114 2188 ± 84 2291 ± 185 2389 ± 129
β-Phenyle hanol 6538 ± 155 6240 ± 257 6571 ± 210 8521 ± 2431 6276 ± 304 5947 ± 347 6607 ± 431 7334 ± 677 3834 ± 148 4351 ± 22 4111 ± 133 4192 ± 320 4861 ± 911 4465 ± 567 4680 ± 938 4894 ± 879
γ-Bu y olac one 173 ± 6 657 ± 56 823 ± 31 1089 ± 204 237 ± 23 716 ± 38 811 ± 54 1076 ± 141 95.2 ± 49.5 959 ± 20 1022 ± 138 1115 ± 9 682 ± 503 819 ± 41 977 ± 65 1149 ± 36
γ-nonalac one 4.2 ± 0.0 4.2 ± 0.0 4.6 ± 0.0 4.5 ± 0.1 4.8 ± 0.1 4.8 ± 0.1 4.7 ± 0.5 5.3 ± 0.2 3.2 ± 0.1 3.1 ± 0.2 3.2 ± 0.2 3.6 ± 0.0 3.6 ± 0.3 3.7 ± 0.0 4.6 ± 0.8 4.2 ± 0.2
γ-decalac one 2.9 ± 0.0 2.9 ± 0.0 3.1 ± 0.1 3.2 ± 0.2 2.6 ± 0.1 2.8 ± 0.1 2.6 ± 0.2 3.0 ± 0.1 1.7 ± 0.2 2.1 ± 0.0 2.2 ± 0.0 1.9 ± 0.2 1.9 ± 0.1 2.1 ± 0.2 2.5 ± 0.6 2.1 ± 0.2
Bu y ic acid 102 ± 39 172 ± 3 89.0 ± 27.3 113 ± 63 116 ± 64 78.2 ± 23.3 168 ± 38 0.0 ± 0.0 494 ± 59 150 ± 21 356 ± 226 160 ± 59 229 ± 114 317 ± 225 242 ± 102 240 ± 104
Isobu y ic acid 479 ± 13 479 ± 13 489 ± 11 413 ± 82 387 ± 59 410 ± 49 409 ± 26 446 ± 13 98.5 ± 44.8 166 ± 2 203 ± 37 227 ± 47 155 ± 6 144 ± 11 140 ± 22 143 ± 18
Hexanoic acid 1007 ± 59 948 ± 4 996 ± 8 1187 ± 263 603 ± 265 578 ± 256 650 ± 279 637 ± 249 488 ± 51 540 ± 54 342 ± 142 556 ± 12 301 ± 8 264 ± 1 291 ± 18 279 ± 14
Oc anoic acid 1824 ± 31 1884 ± 207 1766 ± 172 1680 ± 386 1073 ± 513 1072 ± 535 1183 ± 481 1024 ± 502 984 ± 144 885 ± 55 895 ± 151 1002 ± 95 549 ± 116 505 ± 85 549 ± 60 602 ± 64
Decanoic acid 531 ± 36 500 ± 63 508 ± 98 559 ± 211 364 ± 178 1099 ± 929 509 ± 211 433 ± 197 628 ± 56 434 ± 32 762 ± 361 689 ± 73 814 ± 596 530 ± 318 559 ± 264 954 ± 457
TDN 0.5 ± 0.0 0.2 ± 0.0 0.7 ± 0.0 0.8 ± 0.3 0.5 ± 0.0 15.9 ± 1.2 35.2 ± 3.3 105 ± 5 1.3 ± 0.0 0.1 ± 0.1 0.1 ± 0.1 0.1 ± 0.1 0.5 ± 0.0 10.9 ± 0.7 38.8 ± 11.7 85.4 ± 10.5
β-damascenone 0.2 ± 0.1 0.3 ± 0.0 0.3 ± 0.0 0.1 ± 0.1 2.3 ± 0.3 3.1 ± 0.3 4.0 ± 0.5 4.4 ± 0.2 0.3 ± 0.0 0.1 ± 0.1 0.1 ± 0.1 0.2 ± 0.0 2.3 ± 0.3 3.2 ± 0.1 3.9 ± 0.2 4.0 ± 0.1
β-ionone 0.6 ± 0.0 0.5 ± 0.0 0.3 ± 0.0 0.6 ± 0.3 0.7 ± 0.0 0.5 ± 0.0 0.3 ± 0.0 0.4 ± 0.0 0.4 ± 0.0 0.3 ± 0.1 0.5 ± 0.0 0.4 ± 0.0 0.6 ± 0.0 0.2 ± 0.0 0.2 ± 0.0 0.3 ± 0.0
E hyl cinnama e 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.1 ± 0.0 0.2 ± 0.0 0.2 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.1 ± 0.0 0.2 ± 0.0
Linalool 2.4 ± 0.1 3.0 ± 0.0 1.5 ± 0.0 2.5 ± 1.2 73.8 ± 1.7 40.6 ± 1.1 10.2 ± 0.7 2.1 ± 0.0 3.5 ± 0.0 1.2 ± 0.1 1.7 ± 0.6 1.8 ± 0.1 65.3 ± 1.6 57.3 ± 2.5 11.2 ± 4.5 2.8 ± 0.5
α- e pineol 1.0 ± 0.0 1.6 ± 0.0 1.2 ± 0.0 1.0 ± 0.3 30.8 ± 1.9 138 ± 4 116 ± 3 48.4 ± 3.8 1.1 ± 0.0 1.2 ± 0.2 0.9 ± 0.3 0.7 ± 0.1 26.6 ± 1.6 134 ± 6 121 ± 11 66.8 ± 9.0
β-ci onellol 1.2 ± 0.0 0.8 ± 0.0 0.5 ± 0.0 0.7 ± 0.3 5.6 ± 0.7 2.2 ± 0.2 0.7 ± 0.0 0.6 ± 0.1 1.4 ± 0.1 0.4 ± 0.1 0.3 ± 0.3 0.5 ± 0.1 4.4 ± 0.1 1.7 ± 0.2 0.8 ± 0.1 0.7 ± 0.1
Ge aniol 1.1 ± 0.1 0.0 ± 0.0 0.3 ± 0.1 0.5 ± 0.6 13.3 ± 0.9 12.5 ± 1.4 2.9 ± 0.1 0.0 ± 0.0 1.1 ± 0.0 1.3 ± 1.0 1.4 ± 1.0 0.4 ± 0.1 12.4 ± 0.0 17.8 ± 1.1 3.5 ± 1.2 0.0 ± 0.0
Guaiacol 0.1 ± 0.1 0.1 ± 0.0 0.1 ± 0.0 0.2 ± 0.2 0.9 ± 0.3 0.4 ± 0.0 0.8 ± 0.0 1.1 ± 0.0 0.3 ± 0.1 0.1 ± 0.0 0.1 ± 0.0 0.1 ± 0.0 0.7 ± 0.1 0.7 ± 0.0 0.9 ± 0.3 0.9 ± 0.0
4- inylguaiacol 17.6 ± 5.1 13.0 ± 6.0 23.0 ± 3.0 60.3 ± 35.7 525 ± 156 1114 ± 65 1719 ± 117 2827 ± 25 45.9 ± 15.0 13.9 ± 10.9 28.8 ± 24.8 22.2 ± 1.0 399 ± 58 1055 ± 29 1748 ± 186 2458 ± 56
2-6-dime hoxyphenol 0.9 ± 0.9 0.0 ± 0.0 0.1 ± 0.1 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 1.9 ± 1.9 0.0 ± 0.0 0.3 ± 0.1 0.2 ± 0.0 0.1 ± 0.1 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
E-isoeugenol 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.5 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
4- inylphenol 22.1 ± 2.8 21.4 ± 6.5 15.8 ± 1.6 30.4 ± 23.8 643 ± 230 1094 ± 169 1383 ± 96 2243 ± 41 134 ± 71 75.7 ± 58.4 80.9 ± 57.4 10.2 ± 10.2 469 ± 126 793 ± 82 1463 ± 164 1874 ± 21
anillin 8.1 ± 7.1 1.0 ± 0.1 0.9 ± 0.0 1.6 ± 0.5 5.5 ± 0.6 7.4 ± 0.5 7.4 ± 0.1 16.5 ± 10.0 1.1 ± 0.3 0.7 ± 0.4 0.4 ± 0.4 1.3 ± 0.1 5.4 ± 0.4 7.9 ± 0.5 8.2 ± 0.3 5.2 ± 0.2
ace o anillone 5.8 ± 5.8 0.4 ± 0.4 0.2 ± 0.2 0.6 ± 0.1 49.6 ± 4.5 43.6 ± 3.9 41.5 ± 2.3 36.6 ± 9.7 0.5 ± 0.5 0.0 ± 0.0 0.4 ± 0.4 0.0 ± 0.0 48.8 ± 3.5 47.2 ± 7.1 42.8 ± 3.2 27.8 ± 2.3
PR
L. he mo ole ans
PR
CTL
T. delb ueckii
CTL
S.d. Table 5 (con ) A e age concen a ion o ola iles measu ed abo e he limi o quan i ica ion in con ol wines and hose spiked wi h glycosidic p ecu so s
om Riesling g apes in un e men ed con ols (Acid Hyd olysis) and in wines e men ed wi h S. ce e isiae. P. kluy e i. T. delb ueckii and L. he mo ole ans.
252
S.d. Table 6 A e age alues o compounds abo e he limi o quan i ica ion acco ding he yeas s ain ha
ca ied e men a ion and acidic hyd olysis (AH) o wines wi h o wi hou Riesling p ecu so s. The le e s
a-d exp ess he ducan pos -hoc es being a he highes alue.
AH
P. kluy e i
S. ce e isiae
T. delb ueckii
L. he mo ole ans
e hyl ace a e
0 c
26053 a
17723 b
22287 ab
19027 b
isoamyl ace a e
0 b
310 a
21 b
9.8 b
12 b
isobu yl ace a e
1.6 c
152 a
19 b
17.4 b
8 c
phenyle hyl ace a e
2.3 b
2177 a
16.4 b
32.4 b
5.6 b
e hyl hexanoa e
0 e
151 a
79.7 b
49.4 c
33.5 d
e hyl oc anoa e
0 d
160 a
93.3 b
31 c
26.5 c
e hyl decanoa e
0 c
99 a
51.3 b
40.7 b
13.2 c
e hyl isobu y a e
0 d
29.7 b
14.5 c
37.8 a
13.6 c
e hyl 2-me hylbu y a e
0 d
2.2 a
1.7 b
2.5 a
0.9 c
e hyl lac a e
0 b
590 b
582 b
514 b
2369 a
isobu anol
0 d
8664 a
8534 a
7742 b
4890 c
isoamyl alcohol
0 c
33579 b
43957 a
35517 b
33612 b
me ionol
0 d
4457 a
3344 b
4538 a
1912 c
β-phenyle hanol
0 d
6322 a
5346 b
6754 a
4424 c
e hyl lac a e
0 b
590 b
582 b
514 b
2369 a
γ-bu y olac one
0 c
819 ab
776 ab
698 b
852 a
𝛾-nonalac one
0 d
5.1 a
4.6 b
4.6 b
3.7 c
𝛾-decalac one
0 d
3.8 a
2.8 b
2.9 b
2.1 c
bu y ic acid
0 b
551 a
210 ab
105 b
274 ab
isobu y ic acid
0 d
397 b
165 c
439 a
159 c
hexanoic acid
0 e
2198 a
1137 b
826 c
383 d
oc anoic acid
0 e
3963 a
2510 b
1438 c
746 d
decanoic acid
0 d
1218 a
845 b
563 c
671 bc
TDN
7 a
9.2 a
20.3 a
19.9 a
17.1 a
β-damascenone
3.3 a
1.4 c
1.9 b
1.8 b
1.8 b
linalool
44.5 a
11.8 b
15.7 b
17 b
18.1 b
α- e pineol
70.9 a
24.5 b
39.7 b
42.2 b
44 b
β-ci onellol
0.4 b
1.2 a
1.9 a
1.5 a
1.3 a
Ge aniol
13.1 a
10.003 ab
3.381 c
3.840 c
4.733 bc
4- inylguaiacol
999 ab
722 b
1218 a
787.5 ab
721.2 b
4- inylphenol
876 b
636 b
1496 a
681.8 b
612.3 b
anillin
20.4 a
4.4 b
7.7 b
6.1 b
3.8 b
ace o anillone
4.2 d
15.9 c
29.9 a
22.3 b
20.9 b
Supplemen a y da a
253
4. Supplemen a y da a om Sec ion II: Chap e 2
S.d. Table 7 3-way ANOVA assessing he e ec o he ac o s: p esence o absence o p ecu so s. yeas
s ain. aging and hei in e ac ion on he ola ile composi ion o Ga nacha syn he ic wine.
P > F
P ecu so s
Yeas
Aging
P ecu so s*
Yeas
P ecu so s*
Aging
Yeas *
Aging
E hyl ace a e
< 0.0001
0.00
n.s
n.s
0.02
n.s
Isoamyl ace a e
n.sa
< 0.0001
n.s
n.s
n.s
0.01
E hyl hexanoa e
0.00
< 0.0001
0.00
0.01
0.03
0.02
E hyl oc anoa e
0.00
< 0.0001
0.01
0.01
n.s
n.s
E hyl decanoa e
0.01
0.00
n.s
n.s
n.s
n.s
Isobu anol
n.s
0.01
0.01
n.s
n.s
n.s
1-Bu anol
n.s
0.00
0.05
n.s
n.s
n.s
Isoamyl alcohol
0.01
< 0.0001
n.s
n.s
n.s
n.s
1-Hexanol
< 0.0001
< 0.0001
0.00
0.00
0.01
n.s
Me ionol
0.01
< 0.0001
< 0.0001
n.s
n.s
n.s
β-Phenyle hanol
n.s
< 0.0001
n.s
0.04
n.s
n.s
E hyl lac a e
0.04
< 0.0001
< 0.0001
0.02
n.s
< 0.0001
γ-Bu y olac one
0.00
< 0.0001
< 0.0001
0.01
n.s
< 0.0001
Bu y ic acid
n.s
< 0.0001
0.00
n.s
n.s
0.00
Isobu y ic acid
0.05
< 0.0001
n.s
n.s
n.s
n.s
Hexanoic acid
n.s
< 0.0001
0.01
n.s
n.s
n.s
Oc anoic acid
n.s
n.s
n.s
n.s
n.s
n.s
Decanoic acid
0.01
< 0.0001
0.01
0.00
n.s
0.02
E hyl isobu y a e
0.01
< 0.0001
< 0.0001
n.s
n.s
0.00
Isobu yl ace a e
0.05
< 0.0001
0.00
0.04
n.s
0.01
E hyl 2-me hylbu y a e
0.00
< 0.0001
< 0.0001
0.03
0.05
0.01
E hyl iso ale a e
0.00
< 0.0001
< 0.0001
0.01
0.05
< 0.0001
Phenyle hyl ace a e
0.01
< 0.0001
< 0.0001
0.00
n.s
< 0.0001
γ-nonalac one
0.01
0.01
n.s
n.s
n.s
0.05
γ-decalac one
n.s
< 0.0001
n.s
n.s
n.s
n.s
TDN
< 0.0001
n.s
< 0.0001
n.s
< 0.0001
n.s
β-damascenone
< 0.0001
n.s
0.00
n.s
0.00
n.s
Linalool
< 0.0001
n.s
< 0.0001
n.s
< 0.0001
n.s
α- e pineol
< 0.0001
n.s
< 0.0001
n.s
< 0.0001
n.s

254
P > F
P ecu so s
Yeas
Aging
P ecu so s*
Yeas
P ecu so s*
Aging
Yeas *
Aging
β-ci onellol
< 0.0001
n.s
< 0.0001
n.s
0.03
n.s
Ge aniol
< 0.0001
0.01
0.00
n.s
n.s
0.03
Guaiacol
< 0.0001
0.01
0.00
0.01
0.00
n.s
4- inylguaiacol
< 0.0001
0.04
< 0.0001
0.03
< 0.0001
n.s
2-6-
dime hoxyphenol
< 0.0001
0.02
0.00
0.03
0.00
n.s
E-isoeugenol
< 0.0001
0.00
n.s
0.00
n.s
n.s
4- inylphenol
< 0.0001
n.s
0.00
n.s
0.01
n.s
anillin
< 0.0001
n.s
n.s
n.s
n.s
n.s
ace o anillone
< 0.0001
n.s
n.s
n.s
n.s
n.s
a n.s – no signi ican
Supplemen a y da a
255
S.d. Table 8 A e age concen a ion o ola iles measu ed abo e he limi o quan i ica ion in con ol wines and hose spiked wi h glycosidic p ecu so s om
Ga nacha g apes in un e men ed con ols (Acid Hyd olysis) and in wines e men ed wi h S. ce e isiae. P. kluy e i. T. delb ueckii and L. he mo ole ans.
P ecu so s
Yeas
Aging 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5
E hyl ace a e 0 0 0 0 159 81.4 113 89.8 47916 ± 25983 40643 ± 400 51373 ± 815 75038 ± 3552 98265 ± 815 103863 ± 1277 105766 ± 8826 111643 ± 2094 115396 ± 3465 53785 ± 53547 134323 ± 6716 126562 ± 6215 188802 ± 1965 181048 ± 5838 171344 ± 4821 154900 ± 482
Isoamyl ace a e 0 0 0 0 0 0 0 0 226 ± 107 407 ± 18 325 ± 8 233 ± 5 377 ± 24 377 ± 1 319 ± 27 224 ± 2 3415 ± 147 2998 ± 332 2727 ± 279 1378 ± 17 2997 ± 128 2557 ± 10 2235 ± 100 1191 ± 60
Isobu yl ace a e 0 0 0.66 0.63 7.3 4.2 6.1 5.3 94.6 88.6 ± 5.4 96.2 ± 7.0 87.6 ± 4.6 88.2 ± 0.3 83.1 ± 2.9 82.3 ± 0.7 80.6 ± 1.8 219 ± 12 195 ± 10 186 ± 5 171 ± 13 252 ± 5 205 ± 0 197 ± 3 176 ± 7
Phenyle hyl ace a e 0 0 0 0 0 0 0 0 133 96.9 ± 5.2 150 ± 67 54.5 ± 1.2 150 ± 3 103 ± 0 92.6 ± 4.3 59.9 ± 0.9 9454 ± 676 7253 ± 446 4765 ± 365 9850 ± 94 6837 ± 363 6056 ± 318 3570 ± 354
E hyl hexanoa e 0 0 0 0 0 0 0 0 49.5 ± 0.5 218 ± 20 192 ± 22 544 ± 4 0.0 ± 0.0 628 ± 40 602 ± 54 595 ± 35 165 ± 13 179 ± 12 236 ± 1 210 ± 39 359 ± 54 618 ± 120 535 ± 39 501 ± 2
E hyl oc anoa e 0 0 0 0 0 0 0 0 123 ± 123 294 ± 124 369 ± 27 929 ± 13 382 ± 82 1038 ± 83 1046 ± 66 1028 ± 17 235 ± 38 198 ± 75 408 ± 44 308 ± 86 199 ± 37 694 ± 42 827 ± 47 793 ± 40
E hyl decanoa e 0 0 0 0 0 0 0 0 75.7 ± 75.7 104 ± 51 128 ± 11 234 ± 1 109 ± 9 493 ± 37 424 ± 40 128 ± 128 89.2 ± 11.4 50.6 ± 5.5 57.9 ± 57.9 77.5 ± 32.7 109 ± 13 265 ± 96 317 ± 15 155 ± 17
E hyl isobu y a e 0 0 0 0 0 0 0 0 55.5 146 ± 9 250 ± 29 386 ± 6 30.8 ± 0.5 117 ± 10 157 ± 3 277 ± 0 35.0 ± 5.2 141 ± 3 216 ± 15 400 ± 44 40.2 ± 5.1 112 ± 11 177 ± 21 320 ± 7
E hyl iso ale a e 0 0 0 0 0 0 0 0 6.6 21.1 ± 1.5 34.5 ± 2.4 58.8 ± 4.1 4.7 ± 0.1 16.3 ± 1.1 23.8 ± 0.1 51.6 ± 3.4 4.4 ± 0.1 14.8 ± 0.3 26.0 ± 0.1 54.8 ± 4.5 3.6 ± 0.0 13.1 ± 0.8 20.3 ± 0.3 45.6 ± 1.1
E hyl 2-me hylbu y a e 0 0 0 0 0 0 0 0 5.0 14.9 ± 1.2 24.3 ± 0.1 43.4 ± 2.3 3.6 ± 0.0 11.8 ± 0.8 17.5 ± 0.4 37.5 ± 1.8 4.7 ± 0.3 15.2 ± 0.2 26.4 ± 0.7 53.3 ± 4.2 4.5 ± 0.9 11.1 ± 0.6 17.3 ± 0.6 40.2 ± 0.8
E hyl lac a e 0 0 0 0 0 0 0 0 597 ± 0 1973 ± 3 2929 ± 12 4638 ± 64 524 ± 20 2027 ± 58 2914 ± 42 5187 ± 84 724 ± 26 2895 ± 38 4954 ± 3 7909 ± 163 771 ± 12 3111 ± 55 4739 ± 220 8145 ± 156
Isobu anol 0 0 0 0 0 0 0 0 45325 ± 1531 43427 ± 6037 74678 ± 12438 41391 ± 4333 32287 ± 2788 33469 ± 1743 80738 ± 43781 34710 ± 62 50869 ± 6169 46562 ± 4653 72764 ± 21711 46414 ± 1564 45094 ± 1215 60932 ± 20067 86347 ± 11814 43866 ± 750
1-Bu anol 0 0 0 0 0 0 0 0 282 ± 84 319 ± 28 568 ± 91 307 ± 36 361 ± 43 340 ± 8 811 ± 480 354 ± 34 205 ± 20 184 ± 1 290 ± 95 187 ± 14 186 ± 20 290 ± 102 401 ± 45 220 ± 3
Isoamyl alcohol 0 0 0 0 0 0 0 0 191149 ± 21601 213686 ± 16738 221999 ± 21432 220444 ± 17295 192907 ± 1604 187282 ± 1231 202726 ± 8881 197827 ± 7893 183988 ± 396 180723 ± 8648 188716 ± 4510 185446 ± 9139 182229 ± 2320 178591 ± 5591 184375 ± 10388 198478 ± 892
1-Hexanol 0 0 0 0 0 0 0 0 51.1 ± 2.2 31.0 ± 1.0 29.2 ± 3.6 27.0 ± 2.4 127 ± 14 101 ± 4 97.2 ± 2.4 99.6 ± 5.5 32.5 ± 0.6 28.5 ± 0.3 26.5 ± 0.3 24.9 ± 3.5 79.0 ± 10.3 55.0 ± 0.4 59.3 ± 2.0 57.4 ± 1.5
Me ionol 0 0 0 0 0 0 0 0 6215 ± 63 6312 ± 500 7113 ± 223 7042 ± 105 6453 ± 209 7297 ± 18 7650 ± 703 6612 ± 209 9467 ± 187 9795 ± 574 11083 ± 103 11266 ± 208 10042 ± 207 11590 ± 696 11968 ± 402 12035 ± 82
β-Phenyle hanol 0 0 0 0 0 0 0 0 43093 ± 6427 28107 ± 972 29620 ± 2488 32684 ± 5416 30153 ± 1324 33933 ± 112 36173 ± 4712 34833 ± 2229 30452 ± 1547 26911 ± 655 26205 ± 643 30113 ± 165 27534 ± 803 34197 ± 423 34468 ± 2013 30689 ± 422
γ-Bu y olac one 0 0 0 0 0 0 0 0 1128 ± 192 6520 ± 8 8341 ± 622 9626 ± 48 1384 ± 121 6994 ± 90 8923 ± 562 10098 ± 234 870 ± 38 4026 ± 76 5497 ± 197 6007 ± 123 972 ± 21 4820 ± 357 6143 ± 371 7403 ± 443
γ-nonalac one 0.79 0 0.67 0.56 0.97 0.95 1.43 1.54 7.6 8.1 ± 0.2 14.5 ± 6.9 7.6 ± 0.1 10.0 ± 0.3 9.1 ± 0.4 9.4 ± 0.3 9.0 ± 0.1 8.6 ± 0.1 7.7 ± 0.2 8.2 ± 0.1 8.3 ± 0.5 10.4 ± 0.1 10.0 ± 0.1 10.1 ± 0.1 9.9 ± 0.4
γ-decalac one 0 0 0 0 0 0 0 0 4.9 5.1 ± 0.0 9.5 ± 4.9 4.6 ± 0.1 5.1 ± 0.4 4.3 ± 0.1 4.1 ± 0.1 4.3 ± 0.0 6.7 ± 0.1 6.2 ± 0.2 6.2 ± 0.0 6.3 ± 0.4 6.8 ± 0.1 6.2 ± 0.0 6.0 ± 0.4 5.8 ± 0.0
Bu y ic acid 0 0 0 0 0 0 0 0 652 ± 107 520 ± 18 543 ± 70 504 ± 10 660 ± 26 541 ± 2 646 ± 52 675 ± 32 4020 ± 197 2777 ± 215 2718 ± 174 1887 ± 335 3421 ± 103 3180 ± 213 2767 ± 246 1476 ± 109
Isobu y ic acid 0 0 0 0 0 0 0 0 4618 ± 2256 2467 ± 80 2467 ± 200 2548 ± 117 1831 ± 162 1796 ± 11 1902 ± 102 1905 ± 48 2732 ± 127 2480 ± 154 2588 ± 128 2701 ± 194 2113 ± 135 2145 ± 139 2232 ± 53 2032 ± 72
Hexanoic acid 0 0 0 0 0 0 0 0 1257 ± 826 1359 ± 5 1413 ± 86 1044 ± 40 1905 ± 83 1139 ± 26 1227 ± 65 1140 ± 28 1874 ± 31 1382 ± 109 1304 ± 155 1443 ± 170 1652 ± 6 1048 ± 39 1068 ± 51 1042 ± 12
Oc anoic acid 0 0 0 0 0 0 0 0 3121 ± 2071 3658 ± 68 4202 ± 174 3590 ± 26 5384 ± 158 3851 ± 198 4451 ± 643 4107 ± 18 4965 ± 71 4112 ± 152 3942 ± 438 4264 ± 470 4853 ± 176 3847 ± 46 3946 ± 13 3235 ± 16
Decanoic acid 0 0 0 0 0 0 0 0 333 ± 333 752 ± 304 961 ± 82 1191 ± 12 727 ± 188 961 ± 237 991 ± 13 1349 ± 157 676 ± 12 479 ± 1 1146 ± 135 1087 ± 216 655 ± 8 632 ± 130 825 ± 10 908 ± 76
TDN 0 0 0 0 0 0.81 7.5 24 0.5 0.4 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.8 ± 0.0 8.4 ± 0.5 18.8 ± 1.6 75.9 ± 6.5 0.7 ± 0.1 0.3 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.8 ± 0.1 7.1 ± 1.6 19.1 ± 2.1 66.3 ± 4.9
β-damascenone 0 0 0 0 1 3.5 7.1 7.1 0.4 0.2 ± 0.0 0.1 ± 0.1 0.0 ± 0.0 2.1 ± 0.1 3.4 ± 0.1 3.9 ± 0.4 4.5 ± 0.3 0.2 ± 0.0 0.1 ± 0.1 0.0 ± 0.0 2.2 ± 0.1 4.1 ± 0.0 1.4 ± 1.4 4.1 ± 0.1
β-ionone 0 0 0 0 0 0 0 0 0.6 0.6 ± 0.1 0.3 ± 0.3 0.2 ± 0.0 0.5 ± 0.0 0.7 ± 0.1 0.2 ± 0.1 0.2 ± 0.0 0.8 ± 0.0 0.3 ± 0.0 0.5 ± 0.3 0.4 ± 0.3 0.5 ± 0.0 0.6 ± 0.2 0.5 ± 0.2 0.1 ± 0.0
E hyl cinnama e 0 0 0 0 0 0 0 0 0.0 0.0 ± 0.0 8.6 ± 8.6 1.1 ± 0.1 0.0 ± 0.0 0.2 ± 0.0 0.6 ± 0.1 1.6 ± 0.2 0.0 ± 0.0 0.1 ± 0.0 0.5 ± 0.2 0.0 ± 0.0 0.2 ± 0.0 0.4 ± 0.0 0.7 ± 0.4 1.0 ± 0.4
Linalool 0 0 0 0 1.5 9.6 12.7 6.9 3.3 2.3 ± 0.1 1.0 ± 0.2 0.8 ± 0.0 8.2 ± 0.0 18.4 ± 1.2 12.8 ± 0.1 2.8 ± 0.1 2.4 ± 0.1 2.0 ± 0.1 1.7 ± 0.3 0.8 ± 0.5 7.1 ± 0.1 19.9 ± 0.2 13.1 ± 0.3 2.9 ± 0.1
α- e pineol 0 0 0 0 0 7.4 19.5 23.0 1.1 2.1 ± 0.1 2.4 ± 0.1 1.6 ± 0.2 3.0 ± 0.0 20.5 ± 0.8 25.1 ± 2.0 23.8 ± 0.5 0.7 ± 0.1 1.5 ± 0.0 2.0 ± 0.1 1.8 ± 0.1 2.7 ± 0.1 18.1 ± 0.3 25.2 ± 0.3 26.7 ± 0.3
β-ci onellol 0 0 0 0 0 0 0 0 2.6 1.8 ± 0.1 1.1 ± 0.1 0.3 ± 0.0 4.9 ± 0.2 3.5 ± 0.2 2.6 ± 0.1 0.9 ± 0.1 2.2 ± 0.2 1.4 ± 0.1 1.1 ± 0.1 0.4 ± 0.2 3.4 ± 0.1 2.6 ± 0.1 1.9 ± 0.2 0.8 ± 0.0
Ge aniol 1 0 1 1 2 3 5 3 1.8 0.9 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 3.5 ± 0.2 5.2 ± 0.3 3.6 ± 0.2 1.7 ± 0.0 2.5 ± 0.0 1.3 ± 0.1 1.1 ± 0.0 0.8 ± 0.8 4.4 ± 0.2 5.8 ± 0.2 4.2 ± 0.4 2.3 ± 0.2
Guaiacol 0 0 0 0 0 1 1 2 0.2 0.2 ± 0.0 0.2 ± 0.0 0.1 ± 0.0 4.3 ± 0.3 4.2 ± 0.1 5.1 ± 0.9 5.8 ± 0.6 0.1 ± 0.0 0.2 ± 0.0 0.2 ± 0.0 0.3 ± 0.1 3.2 ± 0.5 2.4 ± 0.2 4.0 ± 0.3 6.3 ± 0.5
4- inylguaiacol 12.7 5.6 5.8 16.5 77.6 127 481 649 20.2 16.3 ± 3.1 18.8 ± 0.6 6.2 ± 1.3 720 ± 5 1588 ± 49 2045 ± 236 2516 ± 123 5.7 ± 1.3 13.7 ± 1.8 12.7 ± 4.3 21.9 ± 0.7 512 ± 59 1277 ± 87 1999 ± 5 2731 ± 75
2-6-dime hoxyphenol 2 0 0 0 0 0.9 3.1 7.0 0.2 0.3 ± 0.1 0.5 ± 0.3 1.0 ± 0.8 11.2 ± 2.6 13.8 ± 3.7 18.4 ± 7.7 19.8 ± 1.7 0.1 ± 0.0 0.2 ± 0.0 0.3 ± 0.1 0.8 ± 0.3 8.3 ± 0.9 4.9 ± 1.0 9.9 ± 0.7 20.2 ± 3.4
E-isoeugenol 0 0 0 0 0 0 0.6 0.7 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 2.5 ± 0.0 2.8 ± 0.3 2.7 ± 0.4 2.4 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 2.3 ± 0.3 2.2 ± 0.2 2.8 ± 0.0 2.8 ± 0.1
4- inylphenol 9.7 4.1 5.3 8.9 38.6 123 264 324 21.5 9.7 ± 9.7 19.9 ± 0.7 9.3 ± 1.8 424 ± 5 569 ± 116 598 ± 174 525 ± 5 11.6 ± 2.7 17.8 ± 1.3 18.3 ± 4.6 22.0 ± 0.1 270 ± 36 346 ± 10 477 ± 24 578 ± 26
Vanillin 10.2 0.6 2.0 1.7 6.9 8.9 21.2 36.8 1.5 0.7 ± 0.1 2.3 ± 1.0 5.7 ± 3.7 10.8 ± 0.7 36.6 ± 23.8 21.5 ± 2.4 20.5 ± 1.2 1.0 ± 0.2 0.8 ± 0.1 1.3 ± 0.4 2.6 ± 1.5 9.4 ± 0.5 9.6 ± 0.2 13.9 ± 4.2 21.5 ± 0.6
Ace o anillone 7.5 1.2 1.1 0.9 53.2 4.2 6.4 12.9 2.5 1.2 ± 0.3 1.5 ± 0.8 3.3 ± 3.3 126 ± 45 194 ± 17 186 ± 6 166 ± 5 0.7 ± 0.1 1.4 ± 0.7 1.0 ± 0.3 2.6 ± 0.5 170 ± 10 131 ± 16 145 ± 34 185 ± 4
Con ol
PG
AH
Con ol
S. ce e isiae
PG
Con ol
P. kluy e i
PG
Supplemen a y da a
256
P ecu so s
Yeas
Aging 0 1 2 5 0 1 2 5 0 1 2 5 0 1 2 5
E hyl ace a e 47032 ± 24509 75440 ± 1213 91377 ± 61 92847 ± 577 166213 ± 2999 155637 ± 7184 148887 ± 15598 127574 ± 458 59220 ± 56 66765 ± 299 67184 ± 1116 75513 ± 265 140554 ± 504 135488 ± 307 131983 ± 1955 100561 ± 177
Isoamyl ace a e 295 ± 182 132 ± 1 122 ± 1 109 ± 8 111 ± 18 144 ± 0 137 ± 3 133 ± 2 217 ± 13 181 ± 7 150 ± 0 113 ± 4 233 ± 33 207 ± 0 167 ± 9 1901 ± 1784
Isobu yl ace a e 88.6 ± 1.6 90.5 ± 1.4 87.6 ± 0.3 111 ± 6 140 ± 45 92.5 ± 0.2 98.2 ± 7.8 118 ± 4 66.4 ± 0.6 61.5 ± 2.6 62.2 ± 3.2 57.9 ± 3.2 67.4 ± 0.4 59.9 ± 3.7 60.4 ± 6.9 68.4 ± 5.6
Phenyle hyl ace a e 320 ± 9 327 ± 83 188 ± 1 142 ± 3 398 ± 139 351 ± 158 168 ± 5 116 ± 5 57.0 ± 7.3 41.2 ± 2.2 37.3 ± 2.5 36.0 ± 11.9 48.2 ± 0.3 57.0 ± 17.7 34.8 ± 1.7 32.4 ± 0.7
E hyl hexanoa e 115 ± 115 95.5 ± 6.5 148 ± 25 103 ± 2 0.0 ± 0.0 135 ± 19 137 ± 41 107 ± 24 48.2 ± 4.7 55.6 ± 1.1 25.2 ± 25.2 54.2 ± 0.2 57.2 ± 57.2 159 ± 2 152 ± 33 144 ± 30
E hyl oc anoa e 268 ± 210 167 ± 7 167 ± 19 177 ± 21 63.8 ± 16.8 154 ± 4 157 ± 33 135 ± 1 93.1 ± 53.2 51.2 ± 8.0 51.4 ± 5.4 63.6 ± 8.1 180 ± 132 181 ± 18 134 ± 6 193 ± 45
E hyl decanoa e 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 49.5 ± 4.9 30.8 ± 3.4 34.2 ± 0.6 61.9 ± 38.1 59.3 ± 14.5 137 ± 40 54.8 ± 54.8 118 ± 27
E hyl isobu y a e 114 ± 4 357 ± 15 505 ± 24 1011 ± 149 132 ± 36 296 ± 0 435 ± 4 816 ± 4 47.9 ± 1.6 139 ± 11 175 ± 36 327 ± 21 38.9 ± 2.1 78.6 ± 64.1 154 ± 19 371 ± 88
E hyl iso ale a e 2.2 ± 2.2 10.7 ± 0.4 16.8 ± 0.9 34.8 ± 2.6 5.2 ± 1.6 9.9 ± 0.1 15.6 ± 0.2 32.9 ± 1.4 2.3 ± 0.2 8.1 ± 0.0 13.2 ± 0.0 26.7 ± 1.8 1.7 ± 0.0 7.2 ± 0.2 11.8 ± 0.7 27.4 ± 0.2
E hyl 2-me hylbu y a e 4.6 ± 0.0 14.1 ± 0.8 21.8 ± 1.1 45.6 ± 2.9 5.7 ± 1.6 12.3 ± 0.4 20.7 ± 0.1 41.6 ± 0.9 2.9 ± 0.2 8.5 ± 0.4 14.2 ± 0.0 28.8 ± 0.7 2.1 ± 0.0 9.2 ± 0.1 12.7 ± 0.2 28.6 ± 0.8
E hyl lac a e 545 ± 54 2442 ± 21 3745 ± 28 6199 ± 11 662 ± 74 2531 ± 94 3882 ± 141 6348 ± 83 16111 ± 817 75862 ± 685 115387 ± 5327 186688 ± 7348 11845 ± 769 63983 ± 1630 95974 ± 2571 146020 ± 9999
Isobu anol 46525 ± 2868 75502 ± 20250 42812 ± 1791 45437 ± 1610 47816 ± 456 45798 ± 1892 44777 ± 335 50035 ± 4681 28429 ± 918 46129 ± 19150 59208 ± 32857 45903 ± 20089 24362 ± 3697 28357 ± 3159 24301 ± 1321 28164 ± 1491
1-Bu anol 314 ± 61 391 ± 108 200 ± 5 219 ± 16 300 ± 5 270 ± 8 275 ± 26 319 ± 27 400 ± 9 643 ± 245 862 ± 474 700 ± 299 436 ± 1 448 ± 90 389 ± 3 474 ± 41
Isoamyl alcohol 199904 ± 33337 167528 ± 3780 169586 ± 8688 165996 ± 3231 165229 ± 782 153452 ± 2168 160373 ± 1153 159103 ± 1830 152611 ± 307 149919 ± 4033 156226 ± 7191 152385 ± 2096 145238 ± 6798 140945 ± 8614 147328 ± 9539 146940 ± 3959
1-Hexanol 49.7 ± 15.2 49.0 ± 0.8 49.1 ± 2.6 47.8 ± 4.4 149 ± 27 107 ± 10 108 ± 0 110 ± 2 33.3 ± 0.4 40.0 ± 3.8 52.8 ± 11.5 38.0 ± 10.7 133 ± 1 113 ± 1 120 ± 11 131 ± 3
Me ionol 6110 ± 9 8165 ± 167 8098 ± 203 8572 ± 224 6363 ± 4 8104 ± 292 9006 ± 325 8776 ± 48 4911 ± 411 6167 ± 11 6519 ± 67 5753 ± 56 5324 ± 485 5890 ± 394 6072 ± 49 6112 ± 39
β-Phenyle hanol 39683 ± 6635 57655 ± 5942 52489 ± 1223 59920 ± 6367 35655 ± 1161 45883 ± 2002 47021 ± 1223 45122 ± 1393 28106 ± 1776 25719 ± 68 27636 ± 789 22502 ± 1023 26220 ± 566 31365 ± 1782 30121 ± 2571 30003 ± 393
γ-Bu y olac one 1061 ± 110 3325 ± 133 3987 ± 169 4650 ± 311 919 ± 37 3213 ± 79 4194 ± 144 4543 ± 66 1548 ± 14 5418 ± 513 6681 ± 371 6665 ± 12 1626 ± 40 5445 ± 307 6666 ± 25 7160 ± 127
γ-nonalac one 9.8 ± 0.2 13.1 ± 4.9 8.6 ± 0.3 8.3 ± 0.3 17.7 ± 6.0 20.9 ± 10.8 10.2 ± 0.3 10.4 ± 0.5 8.1 ± 0.1 7.1 ± 0.1 6.7 ± 0.2 9.9 ± 3.1 9.1 ± 0.0 13.5 ± 5.2 7.7 ± 0.2 8.2 ± 0.1
γ-decalac one 8.4 ± 0.3 10.9 ± 3.9 7.0 ± 0.1 7.1 ± 0.3 11.5 ± 3.8 12.7 ± 6.2 6.2 ± 0.1 6.7 ± 0.3 33.9 ± 3.0 29.4 ± 0.4 28.2 ± 0.3 38.1 ± 11.2 31.7 ± 0.2 44.1 ± 17.7 26.4 ± 0.3 27.8 ± 0.1
Bu y ic acid 639 ± 73 397 ± 32 362 ± 35 494 ± 95 540 ± 29 426 ± 68 499 ± 52 445 ± 54 373 ± 12 329 ± 13 381 ± 54 269 ± 15 423 ± 100 275 ± 22 339 ± 24 289 ± 2
Isobu y ic acid 4645 ± 1870 7041 ± 179 6593 ± 288 7413 ± 767 6339 ± 283 5603 ± 340 6046 ± 374 5682 ± 189 2012 ± 55 2195 ± 92 2466 ± 35 2765 ± 193 2098 ± 174 2048 ± 137 2137 ± 221 2298 ± 225
Hexanoic acid 1110 ± 698 263 ± 2 255 ± 6 231 ± 0 395 ± 26 262 ± 7 258 ± 13 253 ± 8 498 ± 43 409 ± 2 407 ± 12 330 ± 26 509 ± 57 274 ± 30 295 ± 31 279 ± 8
Oc anoic acid 2696 ± 1634 922 ± 37 732 ± 7 871 ± 4 888 ± 120 728 ± 123 731 ± 10 672 ± 26 1263 ± 134 1138 ± 35 1093 ± 64 909 ± 112 1227 ± 80 886 ± 56 827 ± 29 12884 ± 12019
Decanoic acid 419 ± 371 96.2 ± 1.5 98.3 ± 2.3 151 ± 11 19.9 ± 19.9 105 ± 0 110 ± 0 161 ± 16 505 ± 31 585 ± 99 563 ± 26 736 ± 120 1096 ± 115 1316 ± 393 824 ± 39 1254 ± 318
TDN 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.1 ± 0.1 12.8 ± 5.3 21.4 ± 0.9 70.7 ± 10.9 0.1 ± 0.1 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.2 ± 0.0 20.7 ± 6.2 39.3 ± 0.4 136 ± 9
β-damascenone 0.2 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.9 ± 0.6 4.6 ± 2.1 2.9 ± 0.2 3.7 ± 0.1 0.3 ± 0.1 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 2.1 ± 0.3 4.7 ± 1.4 3.9 ± 0.4 4.6 ± 0.1
β-ionone 0.6 ± 0.0 0.2 ± 0.1 0.2 ± 0.0 0.1 ± 0.1 0.8 ± 0.2 0.3 ± 0.3 0.2 ± 0.0 0.1 ± 0.0 0.8 ± 0.1 0.5 ± 0.3 0.1 ± 0.1 0.2 ± 0.0 0.4 ± 0.0 0.1 ± 0.1 0.2 ± 0.0 0.1 ± 0.0
E hyl cinnama e 0.0 ± 0.0 1.2 ± 0.7 0.6 ± 0.1 0.0 ± 0.0 0.1 ± 0.0 3.0 ± 2.7 0.7 ± 0.2 1.2 ± 0.4 0.0 ± 0.0 0.1 ± 0.1 1.6 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 2.2 ± 2.0 1.0 ± 0.4 1.0 ± 0.3
Linalool 3.3 ± 0.0 3.6 ± 0.4 2.3 ± 0.1 0.8 ± 0.1 13.9 ± 4.6 20.2 ± 0.4 11.9 ± 0.5 2.9 ± 0.1 3.4 ± 0.1 2.6 ± 0.2 1.3 ± 0.1 0.5 ± 0.1 10.1 ± 0.3 15.3 ± 0.1 8.3 ± 1.3 1.7 ± 0.0
α- e pineol 1.2 ± 0.0 3.0 ± 0.2 3.5 ± 0.1 3.4 ± 0.0 4.7 ± 1.6 22.7 ± 1.3 26.9 ± 0.8 25.5 ± 0.1 1.2 ± 0.1 2.6 ± 0.5 3.0 ± 0.1 1.8 ± 0.1 3.8 ± 0.0 26.6 ± 2.6 27.6 ± 1.0 21.7 ± 0.2
β-ci onellol 2.7 ± 0.1 2.0 ± 0.4 1.2 ± 0.1 0.3 ± 0.3 6.7 ± 2.3 3.4 ± 0.0 2.2 ± 0.0 0.9 ± 0.1 3.3 ± 0.3 1.9 ± 0.0 1.2 ± 0.0 0.2 ± 0.2 4.6 ± 0.1 3.1 ± 0.3 2.2 ± 0.3 0.7 ± 0.0
Ge aniol 5.9 ± 0.5 2.2 ± 0.7 1.3 ± 0.4 0.6 ± 0.6 13.3 ± 4.4 7.0 ± 0.9 3.7 ± 0.7 1.4 ± 0.4 3.9 ± 0.0 1.2 ± 0.1 0.4 ± 0.4 0.0 ± 0.0 6.0 ± 0.4 5.0 ± 0.1 2.9 ± 0.3 0.5 ± 0.5
Guaiacol 0.2 ± 0.0 0.1 ± 0.0 0.1 ± 0.0 0.2 ± 0.1 3.2 ± 0.4 1.9 ± 0.0 3.0 ± 0.5 4.4 ± 0.2 0.2 ± 0.0 0.2 ± 0.1 0.7 ± 0.1 0.2 ± 0.0 2.2 ± 0.7 2.8 ± 0.1 3.6 ± 0.9 5.6 ± 0.2
4- inylguaiacol 19.2 ± 11.8 13.0 ± 2.4 10.0 ± 2.1 14.9 ± 5.3 787 ± 188 1453 ± 208 2133 ± 204 2657 ± 13 17.5 ± 0.3 19.9 ± 3.8 48.4 ± 25.2 10.7 ± 2.7 582 ± 5 1305 ± 32 1644 ± 54 2113 ± 120
2-6-dime hoxyphenol 0.2 ± 0.2 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 9.6 ± 1.1 5.0 ± 0.8 10.6 ± 3.1 15.9 ± 0.1 0.8 ± 0.4 0.4 ± 0.4 2.4 ± 0.0 0.0 ± 0.0 8.3 ± 1.6 6.6 ± 1.0 13.5 ± 5.5 19.0 ± 0.3
E-isoeugenol 0.2 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 4.1 ± 1.3 3.1 ± 0.4 3.1 ± 0.2 3.2 ± 0.1 0.1 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 1.4 ± 0.0 1.8 ± 0.1 1.6 ± 0.1 2.0 ± 0.2
4- inylphenol 23.0 ± 3.4 18.6 ± 2.9 11.5 ± 1.0 0.0 ± 0.0 403 ± 149 447 ± 128 770 ± 290 560 ± 17 20.5 ± 4.3 25.5 ± 7.9 206 ± 119 0.0 ± 0.0 343 ± 7 331 ± 5 698 ± 277 491 ± 12
Vanillin 1.0 ± 0.3 0.5 ± 0.3 3.2 ± 1.6 2.0 ± 0.0 13.0 ± 4.1 9.3 ± 0.8 25.8 ± 3.2 11.5 ± 11.5 2.7 ± 1.7 1.2 ± 0.3 1.1 ± 1.1 1.2 ± 0.6 22.4 ± 2.1 8.1 ± 0.3 15.4 ± 2.3 20.1 ± 3.2
Ace o anillone 1.5 ± 0.9 0.0 ± 0.0 2.0 ± 0.8 1.2 ± 1.2 212 ± 78 71.7 ± 40.1 172 ± 3 102 ± 60 2.9 ± 2.0 1.0 ± 0.1 0.9 ± 0.9 0.8 ± 0.8 163 ± 1 32.6 ± 2.5 165 ± 3 92.6 ± 60.1
Con ol
PG
T. delb ueckii
Con ol
L. he mo ole ans
PG
S.d. Table 8 (con ) A e age concen a ion o ola iles measu ed abo e he limi o quan i ica ion in con ol wines and hose spiked wi h glycosidic p ecu so s
om Ga nacha g apes in un e men ed con ols (Acid Hyd olysis) and in wines e men ed wi h S. ce e isiae. P. kluy e i. T. delb ueckii and L. he mo ole ans.
Supplemen a y da a
257
S.d. Table 9 A e age concen a ion o ola ile compounds acco ding o he yeas s ain o acidic hyd olysis
con ols o wines wi h and wi hou Ga nacha p ecu so s. Le e s a-d a e he esul s o Duncan pos -hoc
es ; compounds wi h di e en le e s indica e signi ican di e ences
AH
P. kluy e i
S. ce e isiae
T. delb ueckii
L. he mo ole ans
E hyl ace a e
55.4 d
144395 a
79313 c
113126 b
97159 bc
Isoamyl ace a e
0 c
2298 a
311 bc
148 bc
396 b
E hyl hexanoa e
0 c
377 a
354 a
105 b
86.9 bc
E hyl oc anoa e
0 d
490 b
651 a
161 c
118 c
E hyl decanoa e
0 c
147 ab
212 a
0 c
68.1 bc
E hyl isobu y a e
0.2 c
201 b
178 b
458 a
166 b
E hyl 2-me hylbu y a e
0 d
24 a
19.8 b
20.8 b
13.4 c
E hyl iso ale a e
0 d
25.5 a
27.2 a
16 b
12.3 c
Isobu yl ace a e
3 e
197 a
87.6 c
103 b
63 d
Phenyle hyl ace a e
0.1 b
6826 a
105 b
251 b
42.8 b
Isobu anol
0 d
57426 a
48253 b
49838 ab
35606 c
1-Bu anol
0 d
251 c
418 b
286 c
544 a
Isoamyl alcohol
0 e
185508 b
203502 a
167646 c
148949 d
1-Hexanol
0.01 d
47.3 c
70.3 b
83.8 a
82.7 a
Me ionol
0 e
11111 a
6837 c
7899 b
5844 d
β-Phenyle hanol
0 d
30017 c
33575 b
47929 a
27709 c
E hyl lac a e
0 b
4646 b
2599 b
3294 b
88984 a
Bu y ic acid
0 d
2604 a
593 b
475 b
335 c
Isobu y ic acid
0 c
2327 b
2442 b
6170 a
2252 b
Hexanoic acid
0 c
1277 a
1311 a
378 b
375 b
Oc anoic acid
0c
4028 a
4046 a
1030 bc
2529 ab
Decanoic acid
0 c
819 a
908 a
145 b
860 a
γ-nonalac one
0.9 c
9.2 b
9.4 b
12.4 a
8.8 b
γ-decalac one
0.1 c
6.2 b
5.2 b
8.8 b
32.4 a
γ-Bu y olac one
0 e
4981 c
6627 a
3236 d
5151 b
TDN
4 a
13.4 a
13.1 a
13.1 a
24.5 a
β-damascenone
2.3 a
1.7 a
1.8 a
1.5 a
2 a
β-ionone
0.2 b
0.4 a
0.4 a
0.3 ab
0.3 ab
E hyl cinnama e
0.1 a
0.4 a
1.5 a
0.8 a
0.7 a
Linalool
3.9 a
6.8 a
6.2 a
7.4 a
5.4 a
α- e pineol
6.3 a
11.1 a
9.9 a
11.4 a
11 a
β-ci onellol
0.1 c
1.7 b
2.2 ab
2.4 a
2.2 ab
Supplemen a y da a
264
Compound 21 A 22 A 30 A 33 34 so2 med A-16 elemen a-16 7 9a-16
Ace a e Es e s
E hyl ace a e 58606 ± 9924 39129 ± 459 78116 ± 8044 52660,7 56444,1 26193 ± 3530 28778 ± 909 23785 ± 171
Isoamyl ace a e 505 ± 12 617 ± 1 623 ± 170 414,1 502,3 414 ± 56 582 ± 19 223 ± 8
Hexyl ace a e 105 ± 1 128 ± 28 109 ± 9 130,6 137,3 105 ± 27 69.4 ± 5.9 29.0 ± 3.8
Isobu yl ace a e 11.3 ± 0.9 10.3 ± 0.6 10.3 ± 0.3 12,3 11,1 11.1 ± 5.6 8.2 ± 0.0 3.8 ± 0.2
Bu yl ace a e 3.4 ± 1.3 3.2 ± 0.4 4.7 ± 0.1 3,5 4,4 1.0 ± 1.5 2.1 ± 0.8 1.7 ± 0.1
Phenyle hyl ace a e 33.0 ± 1.9 39.7 ± 0.9 39.5 ± 8.2 33,1 38,9 35.9 ± 4.8 90.9 ± 2.5 34.5 ± 0.2
E hyl es e s
E hyl p opanoa e 69.4 ± 10.8 0.0 ± 0.0 26.0 ± 26.0 0,0 78,2 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
E hyl bu y a e 177 ± 16 143 ± 3 226 ± 18 152,0 156,8 81.4 ± 9.8 60.3 ± 16.3 82.4 ± 16.2
E hyl hexanoa e 1481 ± 626 565 ± 18 640 ± 59 668,3 759,4 571 ± 105 428 ± 26 570 ± 18
E hyl oc anoa e 1493 ± 274 749 ± 27 615 ± 135 883,6 884,4 729 ± 167 671 ± 1 529 ± 142
E hyl decanoa e 165 ± 29 98.4 ± 1.9 51.0 ± 16.6 119,8 114,5 75.5 ± 16.0 75.0 ± 1.5 67.3 ± 11.1
E hyl isobu y a e 57.1 ± 4.8 121 ± 4 38.6 ± 2.5 23,6 24,8 25.0 ± 19.6 46.3 ± 1.9 44.5 ± 2.5
E hyl 2-me hylbu y a e 1.6 ± 0.2 2.7 ± 0.1 1.4 ± 0.1 0,9 0,9 1.4 ± 0.6 2.7 ± 1.2 4.6 ± 0.4
E hyl iso ale a e 5.4 ± 0.6 7.1 ± 0.1 4.8 ± 0.2 2,2 2,6 3.2 ± 1.7 11.0 ± 1.2 9.6 ± 1.5
Miscelaneous es e s
E hyl lac a e 6197 ± 66 6297 ± 149 3229 ± 447 13302,8 12613,4 4579 ± 819 4584 ± 137 4980 ± 238
Die hyl succina e 910 ± 37 834 ± 49 364 ± 14 329,8 385,2 466 ± 147 539 ± 38 501 ± 45
Fusel alcohols
Isobu anol 14485 ± 47 25041 ± 186 20180 ± 280 9344,0 9874,8 28096 ± 12995 27662 ± 1455 21434 ± 2448
1-Bu anol 410 ± 1 227 ± 11 586 ± 83 597,8 674,6 519 ± 111 366 ± 32 412 ± 58
Isoamyl alcohol 76299 ± 2091 128039 ± 4359 118899 ± 4508 79937,8 85136,5 124684 ± 20513 212095 ± 8332 170652 ± 2664
1-Hexanol 1254 ± 13 1274 ± 64 1367 ± 96 1014,2 1094,3 1032 ± 15 1222 ± 27 1385 ± 15
c-3-Hexenol 74.9 ± 2.1 75.1 ± 1.5 74.6 ± 1.1 65,3 71,0 7.9 ± 11.2 0.0 ± 0.0 0.0 ± 0.0
Me ionol 455 ± 17 736 ± 28 200 ± 15 154,8 164,5 1293 ± 475 1622 ± 51 1109 ± 36
Benzylic alcohol 92.8 ± 19.2 569 ± 29 44.1 ± 1.6 27,4 37,0 28.9 ± 4.2 19.7 ± 3.5 27.1 ± 0.4
β-Phenyle hanol 10059 ± 1719 13831 ± 92 8343 ± 103 7792,8 8373,5 19778 ± 2028 46768 ± 699 37981 ± 60
Acids
Bu y ic acid 723 ± 6 545 ± 15 1041 ± 42 651,9 709,9 637 ± 65 575 ± 17 659 ± 3
Isobu y ic acid 608 ± 19 1126 ± 25 550 ± 15 336,4 332,9 513 ± 327 860 ± 72 912 ± 65
Iso ale ianic acid 421 ± 23 639 ± 25 440 ± 4 266,4 256,4 395 ± 83 1025 ± 11 1225 ± 1
Hexanoic acid 4515 ± 1024 4640 ± 197 4886 ± 117 5313,6 5712,8 3742 ± 138 2235 ± 60 2686 ± 36
Oc anoic acid 12274 ± 193 11147 ± 22 8163 ± 366 13132,4 12382,0 11386 ± 810 7258 ± 174 8808 ± 181
Decanoic acid 2833 ± 370 2668 ± 225 1369 ± 258 2871,4 3068,8 1846 ± 516 1083 ± 81 1445 ± 53
Mono e penes
Linalool 84.5 ± 4.1 74.2 ± 3.5 67.4 ± 1.2 71,5 76,9 74.4 ± 10.1 79.6 ± 0.4 81.0 ± 1.1
Linalool ace a e 0.7 ± 0.5 0.8 ± 0.2 0.9 ± 0.2 0,5 0,8 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
α-Te pineol 53.3 ± 2.7 50.6 ± 2.0 70.6 ± 1.9 53,6 60,2 63.5 ± 9.3 40.4 ± 0.4 40.0 ± 2.0
β-Ci onelol 3.3 ± 1.1 1.4 ± 1.4 6.4 ± 1.1 4,8 0,0 0.4 ± 0.6 0.6 ± 0.6 0.0 ± 0.0
Ge aniol 11.1 ± 0.1 9.9 ± 0.0 6.7 ± 0.9 10,0 9,4 5.6 ± 2.4 8.9 ± 0.6 10.3 ± 1.0
No isop enoids
β-Damascenone 12.7 ± 1.5 13.6 ± 1.0 9.9 ± 0.6 15,6 13,2 1.2 ± 0.4 2.0 ± 1.0 2.5 ± 1.1
α-Ionone 0.4 ± 0.3 0.7 ± 0.0 0.5 ± 0.0 0,6 0,6 0.0 ± 0.0 0.0 ± 0.0 0.3 ± 0.3
β-Ionone 0.2 ± 0.2 0.3 ± 0.1 0.4 ± 0.1 0,3 0,3 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
Phenols
Guaiacol 5.6 ± 1.0 4.9 ± 1.2 5.3 ± 1.3 5,0 5,0 1.0 ± 0.3 1.0 ± 0.1 1.1 ± 0.6
o-C esol 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,0 0,0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
4-E hylguaiacol 0.3 ± 0.0 0.6 ± 0.2 0.5 ± 0.1 0,2 0,4 0.2 ± 0.2 0.1 ± 0.1 0.1 ± 0.1
m-C esol 0.3 ± 0.0 0.2 ± 0.0 0.2 ± 0.0 0,2 0,2 0.2 ± 0.2 0.0 ± 0.0 0.0 ± 0.0
Eugenol 0.2 ± 0.2 0.0 ± 0.0 0.6 ± 0.0 0,4 0,5 0.5 ± 0.1 0.4 ± 0.2 0.6 ± 0.3
4-E hylphenol 0.4 ± 0.1 0.4 ± 0.1 0.5 ± 0.1 0,3 0,5 0.1 ± 0.2 0.3 ± 0.2 0.3 ± 0.3
4-Vinylguaiacol 102 ± 6 251 ± 3 225 ± 24 130,4 119,3 414 ± 246 278 ± 21 271 ± 46
E-Isoeugenol 2.4 ± 0.2 2.1 ± 0.3 2.0 ± 0.2 2,3 2,3 2.3 ± 1.2 2.6 ± 1.6 3.7 ± 2.1
2,6-Dime hoxyphenol 4.6 ± 1.4 3.1 ± 1.3 3.1 ± 1.2 2,1 2,5 0.3 ± 0.4 0.0 ± 0.0 0.0 ± 0.0
4-Vinylphenol 72.8 ± 1.4 103 ± 4 208 ± 21 69,6 80,8 187 ± 41 123 ± 5 119 ± 1
4-Alyl-2,6-dime hoxyphenol 0.3 ± 0.3 0.0 ± 0.0 0.0 ± 0.0 0,0 0,0 0.1 ± 0.2 0.2 ± 0.2 0.7 ± 0.3
Cinama es
E hyl dihid ocinnama e 0.0 ± 0.0 0.0 ± 0.0 1.1 ± 0.0 0,5 0,8 0.1 ± 0.2 0.8 ± 0.2 1.8 ± 0.0
E hyl cinnama e 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,0 0,0 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
Lac ones
-Whiskylac one 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,0 0,0 0.0 ± 0.0 0.4 ± 0.4 0.0 ± 0.0
c-Whiskylac one 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,0 0,0 2.2 ± 3.1 2.6 ± 2.6 0.0 ± 0.0
γ-Bu y olac one 6337 ± 222 6976 ± 156 3672 ± 143 5552,6 5938,0 3913 ± 712 5854 ± 90 4678 ± 20
γ-Nonalac one 1.4 ± 0.1 1.3 ± 0.1 2.6 ± 0.1 1,1 1,4 0.4 ± 0.6 0.7 ± 0.7 0.5 ± 0.5
γ-Decalac one 17.8 ± 17.8 12.7 ± 1.3 31.0 ± 0.8 19,6 22,4 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
Vanillin de i a es
Vanillin 5.3 ± 1.3 6.5 ± 0.2 7.1 ± 1.4 5,3 6,1 4.7 ± 3.3 5.0 ± 2.9 3.7 ± 2.4
Me hyl anillina e 59.2 ± 4.5 49.1 ± 3.0 59.0 ± 2.8 41,3 47,8 62.5 ± 6.3 59.0 ± 0.2 60.2 ± 0.3
E hyl anilla e 2.0 ± 2.0 1.7 ± 0.2 2.4 ± 0.2 1,7 2,1 0.4 ± 0.1 0.6 ± 0.0 0.5 ± 0.0
Ace o anillone 50.0 ± 6.5 44.8 ± 2.0 54.2 ± 3.3 40,0 44,7 25.0 ± 2.1 24.5 ± 0.9 21.3 ± 2.0
Aldehydes
Benzaldehyde 1.3 ± 0.2 1.7 ± 0.2 2.2 ± 0.2 2,0 1,9 0.2 ± 0.3 0.0 ± 0.0 0.4 ± 0.4
Isobu y aldehyde 10.1 ± 2.1 12.9 ± 0.0 13.5 ± 0.5
Iso ale aldehyde 15.9 ± 2.7 20.1 ± 0.5 18.4 ± 2.6
2-me hylbu anal 1.5 ± 0.2 2.6 ± 0.1 2.3 ± 0.0
Me hional 11.4 ± 2.5 14.5 ± 0.2 12.5 ± 0.2
Phenylace aldehyde 207 ± 11 240 ± 21 248 ± 18
Poly unc ional me cap ans
2-me yl-3- u an hiol 1.2 ± 0.4 1.4 ± 0.7 0.6 ± 0.2 1,362 0,466 0.4 ± 0.3 0.7 ± 0.2 0.6 ± 0.2
Fu u yl hiol 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,005 0,003 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
4-Me cap o-4-me hyl-2-pen anona 0.1 ± 0.1 0.0 ± 0.0 0.0 ± 0.0 0,000 0,000 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
3-me cap ohexyl ace a e 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,000 0,000 0.1 ± 0.1 0.1 ± 0.1 0.0 ± 0.0
3-Me cap ohexanol 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,031 0,029 0.1 ± 0.0 0.1 ± 0.0 0.0 ± 0.0
Benzylme cap an 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0 0,002 0,002 0.0 ± 0.0 0.0 ± 0.0 0.0 ± 0.0
2015
2016
S.d. Table 6 A e age con en o odo an s e men ed in wines om ha es 2015 and 2016wi h di e en
addi i es