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Wine’s Phenolic Compounds and Health:
A Pythagorean View
Francesco Visioli 1,2, * , Stefan-Alexandru Panaite 1 and Joao Tomé-Carneiro 2
1
2
*
Department of Molecular Medicine, University of Padova, Viale G. Colombo 3, 35121 Padova, Italy;
stefan.panaite@outlook.it
IMDEA-Food, CEI UAM + CSIC, 28049 Madrid, Spain; joao.estevao@imdea.org
Correspondence: francesco.visioli@unipd.it
Academic Editors: Paula Silva and Norbert Latruffe
Received: 14 August 2020; Accepted: 5 September 2020; Published: 8 September 2020
Abstract: In support of the J curve that describes the association between wine consumption and
all-cause mortality, researchers and the lay press often advocate the health benefits of (poly)phenol
consumption via red wine intake and cite the vast amount of in vitro literature that would corroborate
the hypothesis. Other researchers dismiss such evidence and call for total abstention. In this review,
we take a skeptical, Pythagorean stance and we critically try to move the debate forward by pointing
the readers to the many pitfalls of red wine (poly)phenol research, which we arbitrarily treat as if they
were pharmacological agents. We conclude that, after 30 years of dedicated research and despite the
considerable expenditure, we still lack solid, “pharmacological”, human evidence to confirm wine
(poly)phenols’ biological actions. Future research will eventually clarify their activities and will back
the current recommendations of responsibly drinking moderate amounts of wine with meals.
Keywords: wine; polyphenols; flavonoids; diet; clinical trials; metabolites
1. Introduction
The association between alcohol consumption and health follows a J-shaped curve [1]. Moderate
alcohol use is associated with better prognosis and lower all-cause death, whereas excessive intake
is detrimental to human health [1]. The mechanisms underlying the protective effects of moderate
alcohol consumption are under investigation and mostly involve reduced plasminogen levels and
lower thrombogenicity observed in moderate drinkers vs. abstainers [1]. Some authors propose the
superiority of wine, namely red wine over other alcoholic beverages and attribute such advantage
to the (poly)phenolic components of red wine [2]. Even though this notion is not fully proven and
is, conversely, often challenged [2], much research is being performed to elucidate the purported
biochemical mechanisms through which wine (poly)phenols would afford better health, in particular
by lowering cardiovascular risk. The debate on alcohol use and health is becoming heavily polarized:
one party underscores a large amount of data in support of the J curve [1,3] whereas the other side
dismisses such evidence and calls for total abstention [2,3]. In support of the former, researchers and
the lay press often advocate the health benefits of (poly)phenol consumption via red wine intake and
cite the vast amount of in vitro literature that would corroborate the hypothesis [4,5].
In this review, we take a skeptical, Pythagorean stance and we critically try to move the debate
forward by pointing the readers to the many pitfalls of red wine (poly)phenol research, which we
arbitrarily treat as if they were pharmacological agents.
2. Phenolic Compounds and Health
Diet and nutrition are essential to promote and maintain good health throughout life and for many
years they have been known to be of crucial importance as risk factors for chronic diseases, making
Molecules 2020, 25, 4105; doi:10.3390/molecules25184105
www.mdpi.com/journal/molecules
Page 2
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)them essential components of prevention activities [6]. The consumption of foods derived from plant
products such as wine, fruits, vegetables, nuts, cereals, legumes, spices and others integrated into
Molecules
2020,
25, PEER
x FORREVIEW
PEER REVIEW
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2020,
25,
x FOR
20 2 of 20
the Mediterranean
or the
DASH
diets
[7], is associated with beneficial health effects and 2aofprotective
role against the development
and
progression
ofpromote
diseases,
as cardiovascular
disease
(CVD)
Dietnutrition
and nutrition
are essential
to
andsuch
maintain
throughout
lifefor
and for [8].
Diet and
are essential
to promote
and maintain
good good
healthhealth
throughout
life and
The ability ofmany
some
plant-derived
foods
to
reduce
disease
risk
has
been
associated
with
the
presence
many
they have
been known
of crucial
importance
risk factors
for chronic
diseases, of
years years
they have
been known
to be to
of be
crucial
importance
as riskasfactors
for chronic
diseases,
making
them
essential
components
of
prevention
activities
[6].
The
consumption
of
foods
derived
them essential
components
of prevention activities
[6].aThe
consumption
of biological
foods derived
non-nutrientmaking
secondary
metabolites
(phytochemicals)
to which
wide
variety of
activities
from
plant
products
such
as wine,
fruits,
vegetables,
nuts,
cereals,
legumes,
spices
and others
from
plant
products
such
as
wine,
fruits,
vegetables,
nuts,
cereals,
legumes,
spices
and
others
are attributed [9,10]. These metabolites have moderate potency as bioactive compounds and low
integrated
intoMediterranean
the Mediterranean
the DASH
dietsis[7],
is associated
with beneficial
integrated
into the
or theorDASH
diets [7],
associated
with beneficial
healthhealth
effectseffects
bioavailability
compared
to
drugs,
but
when
ingested
regularly
and
in
significant
amounts
can have
a protective
role against
the development
and progression
of diseases,
such
as cardiovascular
and aand
protective
role against
the development
and progression
of diseases,
such as
cardiovascular
noticeable mid/long
term
effects.
Phytochemicals
in foods
associated
with a
disease
(CVD)
[8].ability
The ability
of
some
plant-derived
to present
reduce
disease
riskbeen
has
been associated
disease
(CVD)
[8].physiological
The
of some
plant-derived
foodsfoods
to reduce
disease
risk
has
associated
with
theinclude
presence
of non-nutrient
secondary
metabolites
which
a wide
variety
with the
presence
of non-nutrient
secondary
metabolites
(phytochemicals)
to which
a wide
variety
beneficial health
effect
glucosinolates,
terpenoids
and
a(phytochemicals)
large group
oftophenolic
compounds
of biological
activities
are attributed
metabolites
moderate
potency
as bioactive
of biological
activities
are attributed
[9,10].[9,10].
TheseThese
metabolites
have have
moderate
potency
as bioactive
(anthocyanins,
flavones,
flavan-3-ol,
stilbenes,
etc.)
collectively
known
as (poly)phenols
[10,11].
compounds
and bioavailability
low bioavailability
compared
to drugs,
but when
ingested
regularly
and in
compounds
and low
compared
to drugs,
but when
ingested
regularly
and in
significant
amounts
can noticeable
have noticeable
mid/long
term physiological
effects.
Phytochemicals
present
significant
amounts
can have
mid/long
term physiological
effects.
Phytochemicals
present
3. Classification in
and Amounts
of Wine
Phenolic
Compounds
associated
a beneficial
include
glucosinolates,
terpenoids
large group
in foodsfoods
associated
with awith
beneficial
healthhealth
effect effect
include
glucosinolates,
terpenoids
and a and
largea group
of
phenolic
compounds
(anthocyanins,
flavones,
flavan-3-ol,
stilbenes,
etc.)
collectively
as
of compounds
phenolic compounds
(anthocyanins,
flavan-3-ol,
stilbenes,
etc.) collectively
as
Phenolic
have as
a common flavones,
characteristic
in their
chemical
structureknown
the known
presence
of
(poly)phenols
[10,11].
(poly)phenols
[10,11].
one or more hydroxyl groups attached to one or more aromatic or benzene rings. In general, phenolic
3. Classification
and Amounts
of Wine
Phenolic
compounds that
contain
more
than
oneofphenolic
group
areCompounds
called polyphenols to distinguish them from
3. Classification
and Amounts
Wine
Phenolic
Compounds
simple phenolics.
Typically,
thesehave
compounds
arecharacteristic
found
in ainconjugate
form
with
one
or
sugar
Phenolic
compounds
have
as a common
characteristic
in chemical
their chemical
structure
themore
presence
Phenolic
compounds
as
a common
their
structure
the presence
one
or more
hydroxyl
groups
attached
one
more
aromatic
or
rings.
In general,ring
of one
or
more
hydroxyl
groups
attached
to onetoor
more
aromatic
or benzene
rings.to
Inangeneral,
residues linked
byofβ-glycosidic
(O-glycosylated)
bonds
or
byordirect
linkages
ofbenzene
sugar
aromatic
phenolic
compounds
that contain
one phenolic
group
are called
polyphenols
to
compounds
contain
more more
than than
one
group
are called
polyphenols
to
carbon atom phenolic
(C-glycosides)
[12].that
Phenolic
compounds
are phenolic
grouped
according
to their
chemical
structure
distinguish
from simple
phenolics.
Typically,
compounds
are found
in a conjugate
distinguish
them them
from simple
phenolics.
Typically,
these these
compounds
are found
in a conjugate
form form
into two main categories,
flavonoids
and
non-flavonoids,
each comprising
several sub-groups.
In wine,
withorone
or more
residues
by β-glycosidic
(O-glycosylated)
by direct
linkages
with one
more
sugarsugar
residues
linkedlinked
by β-glycosidic
(O-glycosylated)
bondsbonds
or by or
direct
linkages
sub-groups of
flavonoids
compounds
include
flavonols,
flavononols
(also
known
as
dihydroflavonols),
of
sugar
to
an
aromatic
ring
carbon
atom
(C-glycosides)
[12].
Phenolic
compounds
are
grouped
of sugar to an aromatic ring carbon atom (C-glycosides) [12]. Phenolic compounds are grouped
according
to chemical
their
chemical
structure
intomain
twowhile
main
categories,
flavonoids
and non-flavonoids,
according
to their
structure
into
two
categories,
flavonoids
andcontain
non-flavonoids,
each each
anthocyanins,
flavan-3-ols,
flavanones
and
flavones,
non-flavonoids
hydroxycinnamic
comprising
several
sub-groups.
In wine,
sub-groups
of flavonoids
compounds
include
flavonols,
comprising
several
sub-groups.
In
wine,
sub-groups
of
flavonoids
compounds
include
flavonols,
and hydroxybenzoic acids, and stilbenes (Table 1). (Poly)phenolic composition varies among
different
flavononols
(also known
as dihydroflavonols),
anthocyanins,
flavan-3-ols,
flavanones
and flavones,
flavononols
(also known
as dihydroflavonols),
anthocyanins,
flavan-3-ols,
flavanones
and flavones,
wines according while
to thenon-flavonoids
type of grape
used,
vinification
process
used,
type
of
yeast
that
participates
in
contain
hydroxycinnamic
and hydroxybenzoic
and stilbenes
while non-flavonoids contain
hydroxycinnamic
and hydroxybenzoic
acids,acids,
and stilbenes
(Table(Table
1). 1).
the fermentation,
and whether
grape
solids
are
present
in he
maceration
[13].
For
(Poly)phenolic
composition
varies
among
different
wines
according
totype
the of
type
of grape
used,
(Poly)phenolic
composition
varies
among
different
wines
according
to theprocess
grape
used,instance,
vinification
process
used,
type
of yeast
that participates
the fermentation,
andpulp,
whether
grape
solids
vinification
process
used,
type of
yeast
that participates
in thein
fermentation,
andi.e.,
whether
grape
solids
in grapes, the
composition
in
phenolic
compounds
is location-dependent,
skin
and
seeds
are present
the maceration
process
[13].instance,
For instance,
in grapes,
the composition
phenolic
aretypes
present
in proportions
theinmaceration
process
[13]. For
grapes,
theare
composition
have different
and
of
(poly)phenols);
sinceinred
wines
exposed in
to phenolic
allingrape
parts
compounds
is location-dependent,
i.e., pulp,
skinseeds
and seeds
have different
and proportions
of
compounds
is location-dependent,
i.e., pulp,
skin and
have different
types types
and proportions
of
during the vinification
process
they
(poly)phenols
than
white
wines, whose
contents
(poly)phenols);
redcontain
wines
aremore
exposed
all grape
during
the vinification
process
(poly)phenols);
since since
red wines
are exposed
to all to
grape
parts parts
during
the vinification
process
they they
essentially originate
from
the
pulp. than
In this
the
minimum
and essentially
maximum
levels
ofpulp.
total
contain
ore
(poly)phenols
thansense,
white
whose
contents
originate
from
the pulp.
In
contain
more
(poly)phenols
white
wines,wines,
whose
contents
essentially
originate
from the
Inphenolic
this
the minimum
andof
maximum
of phenolic
total phenolic
contents
reported
in a representative
this sense,
minimum
and maximum
levelslevels
of
total
reported
in a representative
contents reported
in sense,
athe
representative
set
studies
(expressed
ascontents
the median
(Q25–Q75)
in mg of gallic
of studies
the median
(Q25–Q75)
inofmg
of gallic
acid equivalents
(GAE)
per liter)
set of set
studies
(expressed
theasmedian
in mg
gallic
acid equivalents
(GAE)
per(89–282)
liter)
acid equivalents
(GAE)
per (expressed
liter)aswere
1531 (Q25–Q75)
(983–1898)
and
3192
(2700–3624),
and
210
and
1531 (983–1898)
and (2700–3624),
3192 (2700–3624),
and(89–282)
210 (89–282)
and(347–434)
402 (347–434)
forand
redwhite
and white
were were
1531 (983–1898)
and 3192
and 210
and 402
for red
402 (347–434) forwines,
red and
white
wines,
respectively
(Table
2).
The
content
of
polyphenols
in
rosé
wine
respectively
2).content
The content
of polyphenols
inwine
rosé wine
is intermediate
between
wines, respectively
(Table(Table
2). The
of polyphenols
in rosé
is intermediate
between
red red
is intermediate
red
and
white wines [14,15].
and white
wines
[14,15].
and between
white
wines
[14,15].
Table
1. Classification
of phenolic
compounds
found
in wine.
1. Classification
of phenolic
compounds
found
in in
wine.
Table 1. Table
Classification
of phenolic
compounds
found
wine.
Group
Subgroup
Subgroup
Group Group Subgroup
Main
Parent
and
Representative
Main Parent
Compounds
and Representative
Derivatives Derivatives
Main Parent
Compounds
and Compounds
Representative
Derivatives
Flavonoids
Flavonoids
Flavonoids
Molecules 2020, 25, x FOR PEER REVIEW
Flavonols
Flavonols
Flavan-3-ols
3 of 20
Kaempferol
Kaempferol
R1 = R2 =R1H= R2 = H
= R2 = OH
Myricetin
Myricetin
R1 = R2 =R1OH
(+)-Catechin R1 = R4 = R5 = OH, R2 = R3 = H
1
Quercetin
R
= H R2 = H
Quercetin R1 = OH, R=2OH,
(-)-Epicatechin R1 = R3 = R5 = OH, R2 = R4 = H
R1
OH
HO
O
HO
HO
HO
O
Molecules
2020,
25, x FOR2 PEER REVIEW
Molecules 2020, 25, x FOR
PEER REVIEW
OH
OH
OH
Procyanidin B1
O
R3
R4
HO
HOH
(+)-Catechin R1 = R4 = R5 = OH, R2 = RHO3 = H
(+)-Catechin R1 = R4 = R5 = OH, R2 = R3 = H OH
Flavan-3-ols
Flavan-3-ols R1 R1
1 = R3 = R5 = OH, R2 = R4 = H
(-)-Epicatechin
R
Cyanidin
R
1
=
OH,
R
2
=
H
OH
(-)-Epicatechin R1 = R3 = R5 = OH, R2 = R4 = H
OH
Anthocyanins
HO
HO
HO
Delphinidin R1 = R2 = OH
HO
O
HO
R2
HO
O
R3=O-glycoside
R2
R3
Malvidin
R1 = R2 = OCH3
R3
R5
R4
R5
R5
R4
HO
Peonidin R1 = OCH3, R2 = H
1 = OH,
R2 = H
OH,
R2 =ROCH
Petunidin R1 =Cyanidin
Cyanidin
R1 3= OH, R2 = H
Anthocyanins
Anthocyanins
Delphinidin R1 = R2 = OH
Delphinidin R1 = R2 = OH
Naringenin R1 = H, R2 = OH
Malvidin R1 = R2 = OCH3
Flavanones
Malvidin R1 = R2 = OCH3
Hesperetin R1 = OH, R2 = OCH3
Peonidin R1 = OCH3, R2 = H
Peonidin R1 = OCH3, R2 = H
HO
O
Flavanones
Flavanones
HO
O
R1
R2
R3
O
OH
OH
HO
O
O
OH
OH
OH
OH
Procyanidin B1
Procyanidin B1
OH
O
Naringenin R1 = H, R2 = OH
Naringenin R1 = H, R2 = OH
Dihydromyricetin R1 = R2 = R3 = OH
OH
OH
R3=O-glycoside
R3=O-glycoside
= OH, R2 = OCH3
Hesperetin
Dihydroquercetin
R1 = R2R=1OH
Hesperetin
R1 = OH, R2 = OCH3
OH
O
Petunidin R1 = OH, R2 = OCH3
Petunidin R1 = OH, R2 = OCH3
Flavanolols
3 of 20
3 of 20
OH
OH
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Molecules
Flavan-3-ols
R1
Flavan-3-ols
Flavan-3-ols
Flavan-3-ols
R1
Flavan-3-ols
Flavan-3-ols
Flavan-3-ols
Flavan-3-ols
R
Flavan-3-ols
Flavan-3-ols
Flavan-3-ols
RR111
HOFlavan-3-ols
O
HO O
HO
HO
O
H
O
HO
O
H
O
H
HHO
O
O
HO O
HO
Group
R1
R11
R
R
OHR
11
OHR 1
OH
OH
OH
O R1 R2
O
O
R2
O
R 3 OO
R2
R
R232OH
R
R33
R
RR433
R
R
33
RR
R
R334
R
R4443
R
R
R
R
44
RR444
RR
42
HO
(+)-Catechin R1 = R4 = R5 = OH, R2 = R3 = H
HO
(+)-Catechin
R
R
R
==2OH,
OH,
R
R
H
(+)-Catechin
R
R
==1111R
==4444OH,
== R
==2222H
(+)-Catechin
R
R
R55555=
OH,
R
R3333==
HHHHOOO
(+)-Catechin
R
==
R
==
R
=R
R
==
R
(+)-Catechin
R1111====R
R4444R
R
OH,
2OH,
R3333R
H
(+)-Catechin
==5555R
== R
R
=R22OH,
== R
R
== H
H
(+)-Catechin
=
=
=
(+)-Catechin
R
=R
=R
=R
(+)-Catechin
R
R
=1R
R
=4OH,
OH,
R
=R
R
=R2H
(+)-Catechin
R5R
=R=OH,
2H
= R33==H
H
2 = R4 = H
(-)-Epicatechin
R1 = R3R=1 =R5R=4 =OH,
R
R
==2OH,
OH,
R
R
H
(-)-Epicatechin
R
R
==1111R
==3333OH,
=R
==2222H
R
OH(-)-Epicatechin
R
R5555=
OH,
R
R4444==
H
(-)-Epicatechin
R
==
R
==
R
=R
R
==
R
(-)-Epicatechin
R
R3333R
R
OH,
2OH,
R4444R
H
(-)-Epicatechin
R1111====R
H
HO
O
==5555R
R
== R
R
=R22OH,
R
== R
R
== H
H
(-)-Epicatechin
R
1R
=
3OH,
=
2H
=
(-)-Epicatechin
OH
OH
===RR
(-)-Epicatechin
R
OH
=R
R
R
=H
(-)-Epicatechin
R
OH
(+)-Catechin
R
1 = RR
4 ==
5=
===ROH,
3 ==
1R
3OH,
= R5R
5R
=2 =OH,
R
2H
= R44==HH
H
(-)-Epicatechin
R
OHHO
OH
O
R2
R22
R
R
R
2
R22
R555
R
R
R
55
R5
R3
Table 1. Cont.
(-)-Epicatechin
R1 = R3 = R5 = OH, R2 = R4 = H
HO
HO
O
HH
O
H
O
O
HO
O
O
OO
HHH
OOO
HO
HHOO
HO
H
HO
O
Anthocyanins
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
Flavanones
OH
OH
OH
OH
OH
OH
OH
OH
3 of 18
OH
OH
OH
OH
OH
OH
HO OH
HHO
OO OH
HHH
OOO OH
HO OH
HO
O
H
HHO
OO OH
HO
O
O
OO
OH
OH
OH
OH B1
OH
OH
OH
Procyanidin
OH OH
OH
OH
OH
Procyanidin
B1
Procyanidin
B1
OH
Procyanidin
B1
Procyanidin
B1
Procyanidin
B1
Procyanidin
B1
OH
Procyanidin
B1
OProcyanidin
B1
Procyanidin
B1
Procyanidin
B1
O
OH
O
OH
O
O
OH
HO O
OH
HO OH
OH
OH
OH
OH
HHOO
Cyanidin R1 = OH, R2 = H
Cyanidin
R
==2OH,
OH,
R
H
Cyanidin
R
OH,
== H
Cyanidin
R11111=
OH,
R22222=
H
Cyanidin
R
=R
R
==
Cyanidin
R1111====OH,
OH,
2OH,
HR
Cyanidin
R
=R22OH,
R
== H
H
Cyanidin
R
HO
Cyanidin
R
=OH,
Cyanidin
R
OH,
R
=H
Cyanidin
R21R
R2 =H
H
== OH
Delphinidin
R1 = R
1OH
=
R
2 = OH
Delphinidin
R
1
=
R
2
=
Delphinidin
R
=R
R2222=
= OH
OH R3=O-glycoside
Delphinidin
R
111OH
=
R
=
OH
Delphinidin
R
1 = R2R
=
Delphinidin
R
=
R
=
OH
Delphinidin
R
1
=
Delphinidin
R
=O-glycoside
== R
Delphinidin
R22==ROH
Delphinidin
R=11OH,
=O-glycoside
Cyanidin
1R
21OH
=H
R2 = OH R
Delphinidin
R33333=O-glycoside
=O-glycoside
R
R33=O-glycoside
=O-glycoside
R
=O-glycoside
1 = R2 = OCH3
Malvidin
RR
R
R
R
33=O-glycoside
=R
=3 OCH
Malvidin
R
R3=O-glycoside
=O-glycoside
R
==1111OCH
Malvidin
R
R22222=
OCH33333
Malvidin
R
=
Malvidin
R2222R
OCH
Malvidin
R1111====R
R
OCH
Malvidin
===2==R
R
===3 OCH
OCH
Malvidin
R
=R
OCH
Malvidin
R
R
=R1R
Malvidin
R
1=
OH
Delphinidin
R
1OCH
R
233= OCH3
Malvidin
Peonidin R1 = OCH3, R2 = H
Peonidin
R
==3,OCH
OCH
3, R2 = H R3=O-glycoside
Peonidin
R
OCH
R
H
Peonidin
R11111=
OCH
3, R
R2222=
H
Peonidin
R
3
Peonidin
R1111====OCH
OCH
R2222====H
H
Peonidin
R
=33,,,OCH
OCH
H
Peonidin
33,,, R
R
=== H
H
Peonidin
R
Peonidin
R
OCH
R
RR
2R
=1=3R
OCH
3H
Malvidin
Peonidin
= 2R
OCH
3 , R2 = H
= OCH
3
Petunidin
RR11==OH,
1R
=
OH,
R
2=
OCH
Petunidin
R
1
=
OH,
2
=
OCH
3
Petunidin
R
= OH,
OH,
R2222=
OCH33333
Petunidin
R1111=
R
Petunidin
R
OH,
2OH,
OCH
Petunidin
R111===OH,
=R
R
==3 OCH
OCH
Petunidin
R
=R
OH,
Petunidin
R
=OH,
Petunidin
R
OH,
==2OCH
OCH
Petunidin
R
Peonidin
1 OCH
3=,22R
=R
H
R2=33=OCH
OCH3
Petunidin
R1R
Naringenin
R11== OH,
H, RR
2 = OH
2==
OCH
Petunidin
R
Naringenin
R
12=
H,
R
2 =3OH
Naringenin
R
1
=
H,
R
OH
Naringenin
RR
= H,
H,
R2222=
OH
Naringenin
R
11=
R
=== OH
Naringenin
R111===H,
H,R
OH
Naringenin
R
H,
R
OH
Naringenin
R
Naringenin
R
R
OH
Naringenin
R
H,
R11212=2=====H,
OH
Naringenin
H,
R2 =OH
OH
2 = OCH3
Hesperetin
R1 = OH,RR
== 2OH,
OH,
R
==3 OCH
OCH
Hesperetin
R
1 = OH,
R
== OCH
Hesperetin
R
OH,
R22222=
OCH33333
Hesperetin
R11111=
=
R
=
Hesperetin
R
1 = OH,
R
2OH,
OCH
3 OCH
Hesperetin
R
=
R
=
Hesperetin
R
OH,
R
Hesperetin
R
OCH
Hesperetin
R
1=
=1 =OH,
OH,
R=222==OH,
=OH
OCH
Hesperetin
R1R
Naringenin
H,RR
1R
R233=OCH
OCH3
Hesperetin
Anthocyanins R4
Anthocyanins
Anthocyanins
Anthocyanins
Anthocyanins
RAnthocyanins
Anthocyanins
Anthocyanins
5
Anthocyanins
Anthocyanins
Anthocyanins
3 of 20
O
O
O
O
O
O
O
OHOH
OH
OH
OH
OH
OH
OH
OH
OH
OH
HO OH
HO
Main Parent Compounds and Representative
Derivatives
Procyanidin
OH B1
OH
HO
O OH
HH
OH
HO
OH
OH
O
OH
O
Subgroup
R
R5
R5
R
RR555
3 of 20
of
20
3 of
20
of 20
20
33
of
of20
203
33 of
of
20
333of
of
20
3 of20
20
OH
OH
OH OH
OH
OH
OH
OH
OH
Hesperetin R1 = OH, R2 = OCH3
R1
Flavanolols
Flavanolols
Flavanolols
FlavanololsRR11
Flavanolols
Flavanolols
Flavanolols
Flavanolols
Flavanolols
Flavanolols
FlavanololsRR11
O
HO
HO O
HOFlavanolols
HO
O O
H
HO
H
O
H
O
OO
HHOO
HO
HO
OH
OH
OH
OH
OH
R2 RR
R1
R
R2RR1111
R
RR222 1
R3
R3
R
RR 3R2
O R
O 1
O
O
O
O
33
OH
OH
OH
OH
OH
O
OH
OH
OH
O OH OH
R3
OH
OH
OH
O
OH
O OH
OH
O
O
O
O
O
OO OH
O
OH
OH
O
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Flavones
Dihydroquercetin R1 = R2 = OH
Dihydroquercetin
R
=R
== OH
OH
Dihydroquercetin
R
R
==1111OH
Dihydroquercetin
R
R22222=
OH
Dihydroquercetin
=
Dihydroquercetin
R1111====R
R2222R
OH
Dihydroquercetin
R
= OH
Dihydroquercetin
R
====R
R
Dihydroquercetin
R
=R
Dihydroquercetin
R
R
=R1OH
Dihydroquercetin
1OH
R2==OH
OH
1 = R2 = R3 = OH
Dihydromyricetin
R
R2
R2
1R
==3R
R
2OH
== R
R
== OH
OH
Dihydromyricetin
RR
1 = R2 R
=
=
Dihydromyricetin
R
R
2
1
R
2
R33333=
OH
Dihydromyricetin
R
1
=
2
=
Dihydromyricetin
R
2
R222R
=1R
=2OH
Dihydromyricetin
R111===R
1R
=333R
R
2OH
=R
R
= OH
Dihydromyricetin
R
=
=
Dihydromyricetin
R2 2
=
=
Dihydromyricetin
R
R
=
R
=
OH
Dihydromyricetin
R
Dihydroquercetin
R
1
=
R
2
=
OH
OH
Dihydromyricetin R1 = R2 = R3==OH
R3
R3
RR
R
3
R333
Dihydromyricetin
R1 = R2 = R3 = OH
OH
OH
OH
OH
OHHO
HO
O
HH
O
H
O
HO
HO
HO
HHOO
OH
HO
HO
OH
OH
OH
OH
OH
OH
OH
O OH
O OH
O
O
O
OH
O OH
O
O
O
O OH
O
O
O O
O
OO H
OO OO O OH O OHOH
HO
O O
O
OH
H
O
OH
OH O
O
O
OH
HHO
OH
O O
OH
OH
O
HOO OO
OH
O
OO CH
O
3
OHO CH
CH3
HO
O CH3
CH33
CH
OHH
H
O
CH
HO CH 3
CH
O33 OH OH CH33
OH HO CH
3
OH
OH
OH O
OH
OH
OH
OH
OH
HO
O
O
O
OO
OH
OH
OH
OH
OH
OH
Dihydroquercetin 3-rhamnosideHHHHOOOO
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
HO
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
Dihydroquercetin
3-rhamnoside
HO
HO
HHO
OO
H
Dihydroquercetin 3-rhamnoside
Apigenin R1 = H
Apigenin
R
=H
Apigenin
R
H
Apigenin
R11111=
H
Apigenin
Apigenin
R1111====H
HR
Apigenin
R
H
Apigenin
R
=== H
H
Apigenin
R
Apigenin
R
H
LuteolinApigenin
R1 = OH R1 = H
1 = OH
Luteolin
R
1
=
OH
Luteolin
R
OH
Luteolin
R111=== OH
Luteolin
R
OH
Luteolin
R111===OH
OH
Luteolin
R
Luteolin
Luteolin
R
Luteolin
R
Apigenin
R1 OH
= RR11==OH
OH
Luteolin
HO
CH3
OH
Luteolin R1 = OH
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Non-flavonoids
Caffeic acid R1 = R4 = H, R2 = R3 = OH
Caffeic
acid
R
=R
42==H,
R
=R
== OH
OH
Caffeic
acid
R
=R
==1111H,
R
==2222OH
Hydroxycinnamic acids
Caffeic
acid
R
RR
4 2=
H,
R
R33333=
OH
=
4
=
CaffeicCaffeic
acidR
R111acid
1acid
R4444R
H,
R
R3333R
OH
Caffeic
R
H,
R
= OH
Caffeic
acid
R
====R
R
4422=
=====H,
H,
R
====R
R
Hydroxycinnamic
acids
Hydroxycinnamic
acids
Caffeic
acid
===R
=R
R
R
=R
Caffeic
acid
R
R
=1H,
R
R
=R2OH
Hydroxycinnamic
acids
Caffeic
1H,
R
4=
H,
2OH
R3==OH
OH
Hydroxycinnamic
Hydroxycinnamic
acids acids
Hydroxycinnamic
acids
Hydroxycinnamic
acids
R4 =RH,
R
2 = OCH3, R3 = OH
Ferulic
acid R1 =acid
Hydroxycinnamic
acids
Hydroxycinnamic
acids
Hydroxycinnamic acids
1H,
=
R
42==H,
R
2=
3, R3 = OH
Ferulic
1acid
=
R
4R
=
R
OCH
3, OCH
R
3 = OH
Ferulic
acid
R
1
=
R
4
=
H,
R
2
=
OCH
R3333=
OH
Ferulic
acid
R
== R
R
44 =
=2===H,
H,
R
=3=3,,OCH
OCH
333,,,, R
R
=== OH
OH
Ferulic
acid
R
1acid
=R
R444R
=11H,
R
OCH
R333===OH
OH
FerulicFerulic
acidR
R11acid
1H,
R
H,
R222=
OCH
R
OH
Ferulic
R
=
4
R
3
=
=
R
2
OCH
R
Ferulic
acid
=
R
=
H,
R
2
=
OCH
3
,
R
OH
Ferulic
acid
R
CaffeicFerulic
acidacid
Racid
1 = R4R
RH,
3R
=2ROH
=2 R
=3 =OCH
=R4R=24==H,
OH3, R3 = OH
p-Coumaric
R1 ==1RH,
Hydroxycinnamic acids
1R
=
R
2H,
=
R
43==H,
R
3 = OH
p-Coumaric
acid
R
1
=
R
2
=
4
=
R
OH
p-Coumaric
acid
R
R
RR4444=
= H,
H,
R3333=
OH
p-Coumaric
acid
R
=
=
R
p-Coumaric
1acid
R222R
=11R
=22H,
OH
p-Coumaric
acidR
Racid
11R
22H,
R
H,
R
OH
p-Coumaric
===4=44R
R
====R
R
R
=== OH
OH
p-Coumaric
R
=R
=R
R
OH
p-Coumaric
acid
==R
R
=R
=R
R3343=3,=====RH,
OH
p-Coumaric
acid
4=
H,
RR
2=
OCH
3=R
OH
Ferulic
acidacid
R1 =RR111acid
1R
2H,
R
H,
3 = OH
p-Coumaric
=acid
OH,
2 = R3 = R4 =
o-Coumaric
==2OH,
OH,
==4R
R
== R
R
== H
H
o-Coumaric
R
1acid
== OH,
R
== R
3R
==2222R
==3333H
o-Coumaric
acid
R
OH,
R
R
R44444=
H
o-Coumaric
acid
R11111=
=
=
o-Coumaric
acid
R
1acid
OH,
R
2OH,
R
3R
R
4R
H
o-Coumaric
acid
R
=
R
=
R
=
R
=
H
o-Coumaric
R
=
OH,
R
2
=
3
=
R
=
o-Coumaric
acid
R
Non-flavonoids
11=
33=
44==
H
o-Coumaric
acid
OH,
R
==R
R
=RRR
=ROH
o-Coumaric
acid R
RR
1==OH,
R2R=1R
R
H,
3=
p-Coumaric
=224=OH,
2R
3H
= R4 =H
H
o-Coumaric
acid
1 = H, R2 = R4 = OCH3, R3 = OH
Sinapic
acid Racid
12==H,
2OCH
== R
R
4=
3, R3 = OH
Sinapic
R
1acid
=
H,
R
R
4R
=
3, OCH
R
3 = OH
Sinapic
acid
R
1
=
H,
R
2
R
4
=
OCH
3
,
R
3
=
OH
Sinapic
acid
R
1
=
H,
R
2
=
4
=
OCH
3
,
R
3
=
Sinapic
acid
R
1
=
H,
R
2
=
R
4
=
OCH
3
,
R
3
=
OH
Sinapic
acid
R
2=
=R
R4 = OCH
R33== OH
OH
Sinapic
acid
RR1122====H,
33,, R
OH
Sinapic
11acid
=acid
=R
R
Sinapic
acid
= H,
H,
R44R
=R2OCH
R343== OH
OH
Sinapic
acid R
R
1R
2OCH
= R434=3=3,,OCH
H
o-Coumaric
acid
RR
=R
OCH
3, R3 = OH
Sinapic
R=1 OH,
2,3-Dihydroxybenzoic acid R1 = R2 = H, R3 = R4 = OH
2,3-Dihydroxybenzoic
acid
R
R
23==H,
R
R
OH
2,3-Dihydroxybenzoic
== R
=3=,1111H,
R,
==3333OH
Hydroxybenzoic acids
= R4 R
=R1acid
OCH
R
=R
Sinapic
acid R1 = H, R2acid
2,3-Dihydroxybenzoic
acid
R
R
H,
R
R4444==
OH
2,3-Dihydroxybenzoic
acid
R
==
R
222OH
=3===R
R
==
R
2,3-Dihydroxybenzoic
acid
1acid
R2222R
H,
R
R4444R
OH
2,3-Dihydroxybenzoic
==3R
R
H,
== R
R
== OH
OH
2,3-Dihydroxybenzoic
=
2
=
Hydroxybenzoic
acids
Hydroxybenzoic
acids
2,3-Dihydroxybenzoic
acid
R
11=
=R
33==
=R
2,3-Dihydroxybenzoic
acid
R
=R
R
=1H,
H,
R
= R,
=R3OH
Hydroxybenzoic
acids
2,3-Dihydroxybenzoic
=R
R
3OH
= R44==OH
OH
Hydroxybenzoic
Hydroxybenzoic
acids acids
Hydroxybenzoic
acids
Hydroxybenzoic
acids
= R3 R
= 1H,
R24==H,
OH
2-Hydroxybenzoic
acid R1 = R2acid
Hydroxybenzoic
acids
Hydroxybenzoic
acids
Hydroxybenzoic
acids
1R
=
R
2H,
=
R
34==H,
R
4 = OH
2-Hydroxybenzoic
acid
R
1
=
R
2
=
3
=
R
OH
2-Hydroxybenzoic
acid
R
=3R
R
=R
RR
==H,
H,
R4444=
= OH
OH
2-Hydroxybenzoic
acid
R
111R
=
R
222H,
=
R
333=
=
H,
R
=
OH
2-Hydroxybenzoic
acid
R
1acid
=
R
2R
=
=
4=
OH
2-Hydroxybenzoic
acid
R
=
R
=
R
H,
R
=
OH
2-Hydroxybenzoic
R
1
=
2
=
3
R
2-Hydroxybenzoic
acid
323=
H,
R
4334=
OH
2-Hydroxybenzoic
acid
R
=R
R212==RR
OH
2-Hydroxybenzoic
acid acid
R11=acid
2,3-Dihydroxybenzoic
=
R
=
R
4=
1R=
R
2H,
=
R
=
H,
R
4OH
=
OH
2-Hydroxybenzoic
4-Hydroxybenzoic acid R1 = R3 = R4 = H, R2 = OH
Hydroxybenzoic acids
==4R
R
== R
R
42==H,
R
== OH
OH
4-Hydroxybenzoic
acid
R
== R
==1111R
==3333H,
OH
4-Hydroxybenzoic
acid
R
R
RR
4 2=
==H,
H,
R22222=
OH
4-Hydroxybenzoic
acid
R
=
=
4
R
4-Hydroxybenzoic
1acid
R3333R
R
4R
H,
R
OH
4-Hydroxybenzoic
acidR
R111acid
R
H,
R
= OH
4-Hydroxybenzoic
==R
==H,
4422=
R
4-Hydroxybenzoic
=1acid
=2R
=3=R
OH
4-Hydroxybenzoic
acid
==R
R
=R=1R
R
OH
4-Hydroxybenzoic
acid
R
RR
=3H,
R4==4===H,
OH
2-Hydroxybenzoic
Racid
1R
=44R
3H,
=RR
R
H,
R2==OH
OH
4-Hydroxybenzoic
3 = OH, R4 = H
Gallic
acid
R1 = R2 = Racid
1R
==3R
R
2OH,
== R
R
3R
==4OH,
OH,
R
4= H
Gallic
1acid
=
R
2R
=
=
=
H
Gallic
acid
R
1
R
2
R
3
OH,
R
4
=
H
Gallic
acid
R
1
=
2
=
3
=
R
4
=
H
Gallic
acid
R
1acid
=R
R222R
=1R
=2OH,
4OH,
=H
HR
Gallicacid
acid
R11acid
1R
=333R
R
2OH,
=R
R33R
=R44OH,
R44== H
H
Gallic
R
=
=
=
Gallic
=
=
=
=
Gallic
R
=
R
=
R
=
OH,
R
=
Gallic
acid
R
R4 R
= 4H,
R2 = OH
4-Hydroxybenzoic
= R2R
= 1R=3 = 3OH,
=H
Gallic
R1acid
1 = R4 = OH, R3 = R2 = H
Gentisic
acid Racid
=R
==3OH,
OH,
R
=R
== H
H
Gentisic
acid
R
== R
==1111OH,
R
== R
==3333H
Gentisic
acid
R
R44444=
OH,
R
R22222=
H
Gentisic
acid
R
=
=
Gentisic
1acid
R4444R
OH,
3OH,
R2222R
H
Gentisic
acidR
R111acid
R
=R33OH,
R
=H
Gentisic
====R
R
=R
R
====R
R
acid
R
=acid
=R
=R
Gentisic
acid
=RR
=R1OH,
=RR
=H
Gentisic
acid
1R=
2R=
31OH,
OH,
4R=
GallicGentisic
acid
R4R
= =OH,
R3H
3H
R2==H
H
Gentisic
1 = R3 = OCH3, R2 = OH, R4 = H
Syringic
acidRR
1OCH
=
R
3=
OCH
3 , R2 R
=
OH,
R
=H
Syringic
acid
R
1
=
R
3
=
3
,
R
2
=
OH,
4
=
H
Syringic
acid
R
=R
R3333=
=3,OCH
OCH
R2222=
= OH,
OH,
R44444=
H
Syringic
acid
R
111OCH
=
333,,,, R
Syringic
acid
1acid
=
R
3R
=
R
2 = OH,
R
4OH,
=
HR
Syringic
acid
R
=
R
=
OCH
R
=
R
H
Syringic
R
1
=
R
=
OCH
3
R
=
OH,
R
=== H
H
Syringic
acid
R
,, R
Syringic
acid
R
==R
RR334==R=OCH
R=22==ROH,
R=44=OH,
=H
Syringic
acid
RR111=acid
OH,
R
3OCH
2OH,
Gentisic
acid
1OCH
=
R
333=
3=, H
R
2R
Syringic
Protocatechuic acid R1 = H, R2 = R3 = OH, R4 = R4 = H
1
=
H,
R
2
=
R
3
=
OH,
R
4= H
Protocatechuic
acid
R
1acid
== H,
R
==H,
R
3R
==222OH,
R
4OH,
== H
Protocatechuic
acid
R
12=
=
H,
=
R
3=
=
R
4=
=H
H
Protocatechuic
R
1
R
=
R
3
OH,
R
4
Protocatechuic
acid
R
1
H,
R
2
R
3
OH,
R
4
H
Protocatechuic
acid
R
1
=
H,
R
=
R
3
=
OH,
R
4
H
Protocatechuic
acid
R
==RR
===44OH,
Protocatechuic
11acid
=acid
H,
R
33R
=OH,
=OH,
Protocatechuic
acid
=OCH
H,RR
R1321,2===R
=H,
=R2OH,
= H RR44===H
Protocatechuic
acid
= RR
3R=
2R=
R33R
4R
H
Syringic
acid
Molecules 2020, 25, x FOR PEER REVIEW
H,
2OH,
H
Protocatechuic
R
R
2 = OH, R3 = OCH3, R4 = H
Vanillic
acid
R1R=1 H,
12==H,
R
2R
==3OH,
OH,
R
3=
3, R4 = H
Vanillic
R
1acid
=
H,
R
OH,
=
OCH
3, OCH
R
4= H
Vanillic
acid
R
1
=
H,
R
2
OH,
R
3
=
OCH
3,
R
4
=
H
Vanillic
acid
R
1
=
H,
R
2
=
R
3
=
OCH
3,
R
4
=
H
Vanillic
acid
R
1acid
=H,
H,R
R
2==H,
OH,
3OH,
=OCH
OCH
R444===H
H
Vanillic
acidR
R11acid
Molecules 2020, 25, x FOR PEER REVIEW
H,
R22R
=R33OH,
R33=3,=3,3,OCH
OCH
3, R
R44== H
H
Vanillic
R
1122=
R
=
R
3,
Vanillic
=
R
=
OH,
=
R
Vanillic
acid
=
H,
R
=
OH,
R
=
OCH
R
H
Vanillic
acid
R
1
=
H,
R
2
=
R
3
=
OH,
R
4
=
H
Protocatechuic
acid
R
Vanillic acid R1 = H, R2 = OH, R3 = OCH3, R4 = H
Resveratrol R1 = R2 = H, R3 = OH
Vanillic acid R1 = H, R2 = OH, R3 = OCH3, R4 = H
Resveratrol R1 = R2 = H, R3 = OH
Piceatannol R1 = H, R3=R2 = OH
Piceatannol R1 = H, R3=R2 = OH
4 of 20
4 of 20
Stilbenes
Stilbenes
Table 2. Range of total phenolic content in red and white wines in a representative set of studies.
Table 2. Range of total phenolic content in red and white wines in a representative set of studies.
Red Wine
Total Phenolic
Content a
Red Wine
Range Content a
Total Phenolic
Min.
RangeMax.
Min.
Max.
1615
4177
1615
1313
1313
2193
4177
2389
2389
3183
2193
622
622
1724
3183
3200
3200
1936
1724
2340
2340
1460
n
7 n
16 7
White Wine
Total Phenolic
White Content
Wine a
Range Content a
Total Phenolic
Min.
RangeMax.
Min.
Max.
216
854
407854
402407
n
7 n
17 7
6 16
20 6
89216
29289
- 292
5 20
282 -
- 402
434 -
1936
3730
3730
3380
23 5
39 23
- 282
210 -
- 434
390 -
1460
2082
2082
1402
3380
3184
3184
3180
3 39
24 3
213210
189213
277390
425277
1402
3200
3200
1788
3180
5900
5900
3070
8 24
- 189
- 425
11 5
- 11
4 8
1788
1012
3070
3264
11 4
55 - 55
370 - 370
20 - 20
4 17
- 4
5 - 5
47 5 47
Reference
Reference
[16]
[16]
[17]
[17]
[18]
[18]
[19]
[19]
[15]
[15]
[20]
[20]
[21]
[21]
[22]
[22]
[14]
[14]
[23]
[23]
Page 4
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)Table 2. Range of total phenolic content in red and white wines in a representative set of studies.
Red Wine
White Wine
Total Phenolic Content a
Total Phenolic Content a
Range
Range
Min.
Max.
n
Min.
Max.
n
Reference
1615
4177
7
216
854
7
[16]
1313
2389
16
89
407
17
[17]
2193
3183
6
292
402
4
[18]
622
3200
20
-
-
-
[19]
1724
1936
5
282
434
5
[15]
2340
3730
23
-
-
-
[20]
1460
3380
39
210
390
47
[21]
2082
3184
3
213
277
5
[22]
1402
3180
24
189
425
11
[14]
3200
5900
8
-
-
-
[23]
1788
3070
4
55
370
20
[24]
1012
3264
11
-
-
-
[25]
554
2669
2
167
347
3
[26]
1181
3589
23
-
-
-
[27]
-
-
-
291
2103
14
[28]
860
2710
8
-
-
-
[29]
1602
1968
7
-
-
-
[30]
1837
3467
6
-
-
-
[31]
38
2
[32]
896
6319
77
83
Mean ± SD
1538 ± 664
3406 ± 1139
189 ± 85
554 ± 545
Median
(Q25–Q75)
1531
(983–1898)
3192
(2700–3624)
210
(89–282)
402
(347–434)
a Data are expressed as gallic acid equivalents (GAE) in mg/L.
3.1. Flavonoids
Flavonoids have a skeleton with 15 carbon atoms and are represented in a C6-C3-C6 type system,
where a benzene ring (designated as B) is joined (in most cases) to the C2 position of a γ-pyran type
ring (C) included in a chromane ring (Table 1) [33]. The structure of flavonoids is shaped by different
levels of hydroxylation, prenylation, alkalization or glycosylation reactions, which give rise to different
sub-groups [34]. In plants, most flavonoids exist as glycosides in combination with monosaccharides
such as glucose and rhamnose (most common), followed by galactose, xylose and arabinose [35].
3.2. Flavonols
Flavonols are characterized by a hydroxyl group in C3 (Table 1) and are often named
3-hydroxyflavones. These compounds are known to play a wide range of biological activities
and are considered the main active compounds within the flavonoids group [36,37]. Flavonols and
their glycosides are present in red and in white wines, influencing their color, taste, and health
properties [38]. Flavonols in red wine include aglycons such as myricetin, quercetin, kaempferol,
and rutin and their respective glycosides (glucosides, glucuronides, galactosides and diglycosides).
Quercetin 3-O-glucoside is the most representative flavonol in wines [39]. Flavonol levels in red wine
can reach over 150 mg/L (Table 3).
Page 5
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)Table 3. (Poly)phenol contents in red and white wines.
Red Wine
Phenol Explorer a
Flavonoids
Main representatives
Anthocyanins
Cyanidin c
Delphinidin c
Malvidin c
Peodinin c
Petunidin c
Total
Dihydroflavonols
Dihydromyricetin
3-O-rhamnoside
White Wine
Phenol Explorer a
USDA b
USDA b
mean
min
max
mean
min
max
mean
min
max
mean
min
max
2.9
16.6
156
18.1
23.6
217
0.6
2.4
12.4
2.5
3.4
21.3
11.9
40.1
541
80.9
61.8
736
1.9
20.1
138
12.5
19.8
193
0.0
0.2
0.0
0.2
0.2
0.6
45.0
57.1
536
50.3
56.6
745
0.4
0.4
0.0
0.0
3.5
3.5
0.6
0.6
0.0
0.0
2.4
2.4
44.7
44.7
44.7
-
-
-
3.0
3.0
3.0
-
-
-
54.4
45.8
59.8
-
-
-
5.7
3.7
15.9
-
-
-
(+)-Catechin
(-)-Epicatechin
Proanthocyanidins
68.1
37.8
355
470
13.8
0.0
99.7
114
390
165
560
1131
71.4
37.9
296
407
0.0
0.0
63.1
63.1
390
165
1354
1917
10.8
9.5
0.2
20.8
0.0
0.0
0.0
0.0
46.0
60.0
1.5
109
7.7
5.5
3.9
17.1
0.0
0.5
0.6
1.1
58.0
60.0
7.3
125
Flavanones
Total
Naringenin c
8.0
8.5
7.3
7.8
8.8
9.4
17.7
24.0
10.3
13.0
25.1
35.0
2.3
2.3
1.7
1.7
2.9
2.9
3.8
7.8
0.0
3.2
7.7
12.5
Flavones
Total
Apigenin
-
-
-
1.3
1.7
0.0
0.0
4.7
8.7
-
-
-
-
-
-
Isorhamnetin c
Kaempferol c
Myricetin c
Quercetin c
5.9
10.2
8.3
44.2
68.6
1.7
5.7
0.0
12.3
19.7
11.6
14.4
17.9
110
154
0.2
0.9
4.2
10.4
15.7
0.0
0.0
0.0
0.0
0.0
1.6
13.7
17.9
33.6
66.8
0.0
0.2
0.0
4.6
4.8
0.0
0.0
0.0
1.3
1.3
0.0
2.6
0.0
20.8
23.4
0.0
0.1
0.1
0.4
0.5
0.0
0.0
0.0
0.0
0.0
0.2
2.7
1.0
8.4
9.4
Total
Flavanols
Total
Flavonols
Total
Page 6
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)Table 3. Cont.
Red Wine
Phenol Explorer a
White Wine
Phenol Explorer a
USDA b
USDA b
mean
min
max
mean
min
max
mean
min
max
mean
min
max
Gallic
Gentisic
Protocatechuic
Syringic
Vanillic
35.9
4.6
1.7
2.7
3.2
70.1
0.0
0.0
0.0
0.0
0.0
13.7
126
8.0
9.6
23.3
7.5
221
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
2.2
18.2
3.3
0.5
0.4
24.8
0.0
0.0
0.1
0.0
0.1
0.4
11.0
20.0
13.0
0.2
1.2
46.8
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Caffeic
Caftaric
Ferulic
(o- and p-) Coumaric
Sinapic
18.8
33.5
0.8
5.8
0.7
100
0.0
1.4
0.0
0.2
0.0
14.9
77.0
179
10.4
40.4
5.4
378
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
2.4
21.5
0.9
1.8
0.6
28.2
0.0
21.4
0.3
0.0
0.0
21.7
7.0
22.0
2.1
5.6
2.8
42.4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Resveratrol d
Resveratrol 3-O-glucoside d
Piceatannol c
Viniferins (δ-. ε-)
Pallidol
5.8
12.5
15.3
7.9
2.0
43.5
0.0
0.0
6.3
0.1
0.0
6.4
61.5
88.0
38.8
26.7
2.5
218
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
0.9
5.0
4.6
0.6
0.7
10.6
0.0
0.4
1.4
0.0
0.0
1.4
3.2
13.1
8.0
0.1
0.3
24.7
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Hydroxybenzaldehydes
Tyrosols
7.1
36.5
43.6
0.0
6.4
6.4
45.6
54.3
99.9
NA
NA
NA
NA
NA
NA
NA
NA
NA
4.1
4.2
8.3
2.4
2.7
5.1
5.8
5.7
11.5
NA
NA
NA
NA
NA
NA
NA
NA
NA
Non-flavonoids
Hydroxybenzoic acids
Total
Hydroxycinnamic acids
Total
Stilbenes
Total
Other polyphenols
Total
a Data are expressed as milligrams of gallic acid equivalent per litter (mg/GAE/L). b Data are expressed as milligrams of gallic acid equivalent per kilogram (mg/GAE/Kg). c Plus glucoside
derivatives.d includes both cis and trans conformations. NA, data not available in the database.
Page 7
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)3.3. Flavan-3-Ols
Flavanols or flavan-3-ols are responsible for the astringency, bitterness, and structure of wines,
and are found in important concentrations in red wine [40]. They are benzopyrans, having no double
bond between C2 and C3, and no C4 carbonyl in Ring C. Furthermore, due to the hydroxylation
at C3 flavanols have two chiral centers. (+)-Catechin (trans configuration) and (−)-epicatechin
(cis configuration) are the two main flavan-3-ol isomers found in red wine, with average combined
concentration over 100 mg/L (Table 3). Catechins usually occur as aglycones, or esterified with gallic acid,
and can form polymers, which are often referred to as proanthocyanidins (or condensed tannins) because
an acid-catalyzed cleavage of the polymeric chains produces anthocyanidins. Proanthocyanidins,
which present an average concentration over 350 mg/L in red wine, include, for example, procyanidin
dimers B1, B2, B3 and B4. Trimers such as procyanidin C1 (three epicatechins) have also been identified.
4. Anthocyanins
Anthocyanic pigments (anthocyanidins and anthocyanins) have a structure based on the
flavylium cation (2-phenylbenzopyrylium). In fact, anthocyanins are anthocyanidin glycosides,
being the corresponding aglycons (anthocyanidins) obtained by hydrolysis. The great variety of
anthocyanins found in nature (more than 500 anthocyanins have been described) is characterized by
the different hydroxylated groups, conjugated sugars and acyl moieties they present [41,42]. The main
anthocyanidins found in red wine are malvidin (most abundant), petunidin, peonidin, delphinidin and
cyanidin. Often anthocyanins are found linked (mainly in position 3) to one or more sugar molecules,
usually glucose, and also to acyl substituents bound to sugars, aliphatic acids, and cinnamic acids.
Anthocyanins can be present in amounts higher than 700 mg/L in red wine, whereas in white wine
they are essentially absent (Table 3).
4.1. Flavanones
Flavanones, also known as dihydroflavones, lack the double bond between carbons 2 and 3 in the
C-ring of the flavonoid skeleton. Some flavanones have unique substitution patterns, e.g., prenylated
flavanones, furanoflavanones, pyranoflavanones, benzylated flavanones, resulting in a large number
of derivatives of this subgroup. One of the main flavonones found in wine is naringenin at levels that
can reach 25 mg/kg (Table 3).
4.2. Flavones
Flavones are characterized by absence of a hydroxyl group in the C3 position and a conjugated
double bond between C2 and C3 in the flavonoid skeleton. Flavones and their 3-hydroxy derivatives
flavonols, including their glycosides, methoxides and other acylated products on all three rings, make
this the largest subgroup among all polyphenols. These compounds were found in grape skin and
wine in both aglycones and glycosides forms. Apigenin, for example, has been described in red wine
only in trace amounts (Table 3).
5. Non-Flavonoids
The non-flavonoid phenolic constituents in wine are divided into hydroxybenzoic acids and
hydroxycinnamic acids, stilbenes and other miscellaneous compounds [43]. These phenolic compounds
can reach levels that range from 60 to 566 mg/L [44].
6. Hydroxycinnamic Acids
Hydroxycinnamic acids are the foremost group of phenolic compounds in grapes and wine [45].
Caffeic, coumaric, and ferulic acids, essentially conjugated with tartaric acid esters or diesters, are some
of the most important compounds in this polyphenol sub-class. For instance, caftaric acid, which is
composed of caffeic acid esterified with tartaric acid, is found in the pulp and represents up to 50% of
Page 8
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)total hydroxycinnamic acids [43,46]. The average amount of hydroxycinnamic acids present in red
wine is around 100 and 30 mg/L in red and white wines, respectively (Table 3).
7. Hydroxybenzoic Acids
In comparison with cinnamic acid derivatives, benzoates are present at lower levels in wine
(Table 3). Hydroxybenzoic acids are phenolic metabolites with a general C6–C1 structure and occur
mainly in their free forms in wine, mainly as p-hydroxybenzoic, gallic, vanillic, gentisic, syringic,
salicylic, and protocatechuic acids [43], although ethyl and mehyl esters of these phenolic acids have
been also identified [47]. Gallic acid, which is present in important levels in white and, especially,
in red wine, is the precursor of all hydrolyzable tannins and is incorporated in condensed tannins [46].
8. Stilbenes
Stilbenes are widely distributed molecules in the Plant Kingdom. However, their presence in the
diet is not very significant, being basically restricted to grapes, red wine and, to a lesser extent, peanuts
and blueberries [48]. Chemically they are 1,2-diarilethenes and usually have two hydroxyl groups in
the meta position of ring A, while ring B is substituted with hydroxyl groups and methoxyl groups in
the meta and/or para positions (Table 1). Although its concentration in wine is much lower than other
polyphenols, i.e., often traces, resveratrol has received much attention for its biological properties and
potential therapeutic effects (see below). The levels of resveratrol aglycone, its piceid glycoside, and its
dimeric and trimeric forms (e.g., pallidol, viniferins) combined may range from negligible up to more
than 100 mg/L (Table 3) when grapes are exposed to fungi.
9. Effects on Human Health
The Greek philosopher Pythagoras of Samos allegedly used to say “All is number” or “God
is number” [49]. He meant that he only believed in what could be measured. This was echoed by
William Thomson, 1st Baron Kelvin who, in his Popular Lectures and Addresses vol. 1 (1889) ‘Electrical
Units of Measurement’, delivered 3 May 1883 notoriously said “When you can measure what you are
speaking about, and express it in numbers, you know something about it, when you cannot express
it in numbers, your knowledge is of a meager and unsatisfactory kind; it may be the beginning of
knowledge, but you have scarcely, in your thoughts advanced to the stage of science.” [50]. What both
scientists meant was that we should base our knowledge on hard evidence. More recently, Dr. Archie
L. Cochrane set out clearly the vital importance of randomized controlled trials (RCTs) in assessing the
effectiveness of treatments [51]. How does this apply to wine (poly)phenols?
We shall start by mentioning that there are thousands of papers published on this topic (a cursory
PubMed search ran on August 14th, 2020 retrieved 2954 entries just by entering “wine polyphenols”).
The near totality of such studies has been performed in vitro. Needless to say, in vitro studies are
indispensable to address mechanisms of action and to propose new avenues of in vivo research.
The case of wine (poly)phenols, however, is rather unique and presents us with a paradigmatic
opportunity to underscore the current very limits of (poly)phenol research.
In keeping with the above, we would like to discuss the case of resveratrol as an example of
molecules for which there exists a strong dyscrasia between the lay public perception of health benefits
and hard scientific data.
Resveratrol became popular in 1991, when Drs. Michel de Lorgeril and Serge Renaud appeared in
the “60 Minutes” CBS show to talk about the French Paradox and to attribute it to the French habit
of drinking red wine, which would theoretically inhibit lipid peroxidation. Note that, back then the
“free radical/antioxidant hypothesis” [52] was in full swing and it was commonplace to believe that
eating and drinking (poly)phenols would scavenge free radicals and prevent their noxious effects,
for example by inhibiting LDL oxidation [53]. This conjecture, now largely proven wrong [54], granted
red wine (poly)phenols, namely resveratrol, immediate popularity and trigger the vast amount of
well-funded research mentioned above.
Page 9
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)Two major issues developed during the nearly three decades that separate the 60 min show from
our current knowledge.
The first problem is that we came to realize that (poly)phenols are very weak (if at all effective)
in vivo direct antioxidants [55]. For kinetic reasons they do not scavenge free radicals and their
bioavailability is generally so low that they contribute very little to the integrated cellular antioxidant
machinery, which is mostly composed of enzymes [56–58]. Alas, plenty of investigators still perform
research and publish papers on the in vitro antioxidant abilities of individual (poly)phenols or of
some raw mixtures of them. Luckily, plenty of researchers correctly use (poly)phenols’ metabolites in
their in vitro studies [59,60]. The hurdle then becomes the difficulty of synthesizing such metabolites,
which are often produced by the organism in different forms. It is worth underscoring that we are
making progress in the identification of metabolites, but—until recently—we mainly focused on
the liver-derived ones [61]. The relatively recent discovery of microbiota-synthesized metabolites
amplifies the list of potential biologically-active molecules produced by the body after the ingestion of
(poly)phenol-rich foods [58,62,63].
In consonance with the above, the lay press often overlooks the bioavailability issue. As regards
resveratrol, already in 1993 Soleas and Goldberg acted as the harbinger of the subsequent in vivo
debacle of the molecule by calling it “a molecule whose time has come and gone” [64]. That conclusive
title might have been a bit too harsh, but it’s a fact that many years of research and many million
dollars invested in it did not yield major results [65,66].
Finally, animal studies often employ very high doses of grape (poly)phenols, e.g., resveratrol and
their results cannot be readily transferred to humans, who would need to ingest several grams of extracts
to replicate the same effects. Indeed, a discrepancy between animal and human effects has just been
underscored [5] and resveratrol’s potential toxicity has been recently reviewed [67]. An often overlooked
paper reported that resveratrol promoted atherosclerotic development in hypercholesterolemic rabbits,
by a mechanism that is independent of observed differences in gross animal health, liver function,
plasma cholesterol concentrations, or LDL oxidative status [68].
10. Human Studies of Resveratrol and Red Wine (Poly)Phenols
One of the fields where red wine (poly)phenols are most actively studied is that of weight control,
namely obesity and its associated insulin sensitivity [69]. The rationale behind studying red wine
(poly)phenols and, particularly, resveratrol is that type II diabetes is rampant in developed countries
and that many researchers are looking for fasting mimetics, to approximate the beneficial effects of
calorie restriction or intermittent fasting on insulin sensitivity [70]. The results are equivocal, to say the
least, as most trials failed to report significant effects, e.g., [71]. The molecular rationale for studying it
is the finding that resveratrol and, maybe, other wine (poly)phenols activate SIRT1, a modulator of
pathways downstream of calorie restriction that produces beneficial effects on glucose homeostasis
and insulin sensitivity [72,73]. This hypothesis is quite controversial for at least two reasons. One is the
factual role of sirtuins as longevity promoters [74]. The other one is that several researchers question
the reproducibility of those data, e.g., [75]. In summary, the jury is still out [76] and the quixotic search
for a substance that would fix the cardiometabolic effects of inordinate diets is not over [77].
Rather than trying to single out individual molecules purportedly responsible for the beneficial
effects of moderate wine use (which is a pharmacological approach), an alternative is to test the effects
of the whole (poly)phenolic fraction. We retrieved 24 publications of human studies that employed
dealcoholized wine (Table S1). Taken together, their results indicate that wine (poly)phenols do exert
healthful effects, ranging from anti-inflammatory actions to modulation of the microbiota, which is now
gaining traction from an industrial viewpoint [78,79] and might be one of the next applications of these
compounds. The extent and precise nature of such activities, however, remains to be fully elucidated.
For example, some publications stem from the same study; there are some contradictions between
data and their discussion (e.g., LPS and LPB data in [80], fatty acid data in [81], inflammatory markers
in [82], etc.); and the true clinical relevance of microbiota modification as related to, e.g., circulating
Page 10
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)lipids (Table S1) [83–106]. In summary, there is indeed evidence that wine (poly)phenols modulate
human physiology, but puffery should be avoided until we can clearly correlate such modifications
with undisputable clinical outcomes.
We also searched the literature for acute or short-term human effects of wine (poly)phenols
(Table S2) [107–123]. Even though this might be seen as a more classic “pharmacological” approach,
even small effects repeated over time might—in the end—affect human physiology and health.
Some outcomes fall in the now-outdated “plasma antioxidant capacity” or “oxLDL” areas, i.e., are
poor proxies of prognosis. Other data are more physiologically relevant and indicate, e.g., salubrious
effects on endothelial function and related flow-mediated dilation. Anti-inflammatory effects have also
been reported. Other studies focused on bioavailability, with scant indications of biological effects. It is
worth noting that ethical reasons often impede research on alcohol in humans [1].
11. Wine vs. Other Alcoholic Beverages: Does Digestion Make the Difference?
Often miscategorized as direct antioxidants [124] (see above) wine (poly)phenols might act as such
during digestion. Several pieces of evidence reveal that, during digestion, lipid peroxides are formed in
the stomach at millimolar concentrations [125]. In addition, we eat pre-formed hydroperoxides, whose
formation is unavoidable in fat-containing foods. Dr. Kanner called the stomach “a bioreactor” [125]
where hydroperoxides are formed and subsequently absorbed. This is particularly noteworthy in the
case of red meat (hypothetically because of its iron content [126]), but it is likely to happen with any
animal food. Lipid peroxidation during digestion can be decreased by the consumption of (poly)phenol
rich foods and beverages such as extra virgin olive oil [127] and—germane to this review—red
wine [128,129]. In a wider context, these data experimentally explain the evolutionary-sound habit of
eating fruit and vegetables, i.e., (poly)phenols with protein [130]. Further, most cultures have culinary
routines that involve drinking (poly)phenols during or after meals, including tea [131], coffee [132,133],
red wine [128], etc.
Another place where wine (poly)phenols might act as indirect antioxidants is the liver, where
ethanol is metabolized to acetaldehyde by the microsomal ethanol oxidizing system (MEOS),
via cyp2E1 [134]. In doing, ROS are generated as by-products. Possibly, (poly)phenols might
lessen this untoward effect of ethanol ingestion, through mechanisms that are yet to be elucidated.
12. Conclusions
In this review, we took a pharma-nutritional approach to wine (poly)phenols. Epidemiological
evidence describes the association between alcohol use and all-cause mortality as following a J-shaped
curve. Many investigators claim the superiority of wine, namely red wine with respect to other
alcoholic beverages and call for (poly)phenols to support their hypothesis. The result is that the lay
public often believes in this premise, in part because there is plenty of in vitro and some animal data
and in part due to wish bias [135]. Indeed, there appears to be a discrepancy between the strength of
biochemical data and the scantiness of well-controlled human trials of individual molecules isolated
from wine. This is commonplace in nutritional research [136,137] and there is no easy way out of
it [138]. In a way, wine (poly)phenols are paradigmatic of the current tension between treating such
compounds as non-essential nutritional agents and expecting pharmacological actions from them [139].
In the former case, we must accept the fact that the biological effects of wine (poly)phenols are minimal
and very difficult to detect with current technologies and biomarkers [138,139]. The latter scenario
involves the unavoidable acceptance of side effects and is not epistemologically applicable to human
nutrition. Another limitation of wine (poly)phenol research is that we often use a reductionist approach
and look for one single mechanism of action. In the case of wine (poly)phenols and particularly
resveratrol, this involves their misclassification as in vivo free radical scavengers and antioxidants,
even if their mechanisms of action are manifold and chiefly involve anti-inflammatory actions and,
possibly, activation of nrf2 and its downstream pathways via xeno-hormesis [1]. In pharma-nutritional
research we should look at a wider picture and acknowledge that phytochemicals contribute to health
Page 11
Voir dans le PDF(s’ouvre dans une nouvelle fenêtre)5 of 20
even though, based on the definition of nutrients, they are not essential. Therefore, these molecules
targets depending on their concentration, and do not have a univocal pharmacological mechanism of
do not fit in the classic and rigorous pharmacological definitions; they can be modified by organisms
action.
before they interact with targets, can have different targets depending on their concentration, and do
In conclusion, after 30 years of dedicated research and despite the considerable expenditure, we
not have a univocal pharmacological mechanism of action.
still lack solid, “pharmacological” human evidence to confirm wine (poly)phenols’ biological actions
In conclusion, after 30 years of dedicated research and despite the considerable expenditure,
(Figure 1). Future research [138] will eventually clarify their activities and will back the current
we still lack solid, “pharmacological” human evidence to confirm wine (poly)phenols’ biological
recommendations of responsibly drinking moderate amounts of wine with meals.
actions (Figure 1). Future research [138] will eventually clarify their activities and will back the current
recommendations of responsibly drinking moderate amounts of wine with meals.
Figure 1. Schematic overview of current pitfalls in wine (poly)phenols research.
Figure 1. Schematic overview of current pitfalls in wine (poly)phenols research.
Supplementary Materials: The following are available online, Table S1: Human long-term studies with
Supplementary
The
are or
available
online,
Table
Human long-term
dealcoholized redMaterials:
wine, Table
S2: following
Human acute
short-term
studies
withS1:
dealcoholized
red wine.studies with
dealcoholized red wine, Table S2: Human acute or short-term studies with dealcoholized red wine.
Author Contributions: This is an Invited Review and all authors (F.V., S.-A.P., and J.T.-C.) contributed to drafting,
revising, Contribution:
and finalizing it.
All is
authors
have read
andand
agreed
the published
versionand
of the
manuscript.
Authors
This
an Invited
Review
all to
authors
(F.V., S.-A.P.,
J.T.-C.)
contributed to
drafting,
revising,
and
finalizing
it.
All
authors
have
read
and
agreed
to
the
published
version
of
the
manuscript.
Funding: This research received no external funding.
Acknowledgments:
This
publication
was not
funded. We thank Paula Silva for inviting us to contribute to this
Funding:
This research
received
no external
funding.
Special Issue. This manuscript forms part of S-A.P. dissertation to obtain his M.D. degree.
Acknowledgments: This publication was not funded. We thank Paula Silva for inviting us to contribute to this
Conflicts of Interest: The authors declare no conflict of interest.
Special Issue. This manuscript forms part of S-A.P. dissertation to obtain his M.D. degree.
Conflicts
of Interest: The authors declare no conflict of interest.
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