RESEARCH ARTICLE
Pos p andial glycaemic esponse o be y nec a s con aining
in e ed suc ose
Rii a Tö önen
1
*, Ja kko Hells öm
2
, Pi jo Ma ila
2
and Kyllikki Kilpi
3
1
Depa men o Clinical Nu i ion, Ins i u e o Public Heal h and Clinical Nu i ion, Uni e si y o Eas e n Finland, PO Box 1627, FI-70211
Kuopio, Finland
2
New Business Oppo uni ies, Na u al Resou ces Ins i u e Finland, Mylly ie 1, FI-31600 Jokioinen, Finland
3
Finnsuga L d, Soke i eh aan ie 20, FI-02460 Kan ik, Finland
(Recei ed 16 May 2016 –Final e ision ecei ed 29 Sep embe 2016 –Accep ed 28 No embe 2016)
Jou nal o Nu i ional Science (2017), ol. 6, e4, page 1 o 7 doi:10.1017/jns.2016.44
Abs ac
Suc ose is commonly used o swee ening be y p oduc s. Du ing p ocessing and s o age o be y p oduc s con aining added suc ose, suc ose is in e ed o
glucose and uc ose. We ha e p e iously shown ha pos p andial glycaemic esponse induced by in ac suc ose is a enua ed when suc ose is consumed
wi h be ies ich in polyphenols. I is no known how in e sion o suc ose a ec s glycaemic esponse. We in es iga ed pos p andial glycaemic and insu-
linaemic esponses o blackcu an (Ribes nig um) and lingonbe y (Vaccinium i is-idaea) nec a s and a e e ence d ink (wa e ) swee ened wi h glucose and
uc ose, ep esen ing comple ely in e ed suc ose. The nec a s and e e ence d ink (300 ml) con ained 17·5 g glucose and 17·5 g uc ose. Polyphenol
composi ion o he nec a s was analysed. A o al o eigh een heal hy olun ee s pa icipa ed in a andomised, con olled, c oss-o e s udy. Blood samples
we e collec ed a as ing and six imes pos p andially du ing 120 min. In e ed suc ose in he e e ence d ink induced glycaemic and insulinaemic esponses
simila o hose p e iously obse ed o in ac suc ose. In compa ison wi h he e e ence, he blackcu an nec a a enua ed he ea ly glycaemic esponse
and imp o ed glycaemic p ofile, and he lingonbe y nec a educed he insulinaemic esponse. The esponses induced by in e ed suc ose in he be y
nec a s a e simila o hose p e iously obse ed o be y nec a s con aining in ac suc ose, sugges ing ha in e sion has no majo impac on glycaemic
esponse o suc ose-swee ened be y p oduc s. The a enua ed glycaemic esponse a e he blackcu an nec a may be explained by inhibi ion o in es inal
abso p ion o glucose by blackcu an an hocyanins.
Key wo ds: Be ies: In e ed suc ose: Pos p andial glucose: Insulin: Polyphenols
Be ies con ain an ioxidan i amins, mine als and fib e, and
hey a e unique as ich sou ces o fla onoids and o he poly-
phenols. Be ies and hei fla onoids, especially an hocyanins,
ha e been associa ed wi h educed isk o CVD
(1–4)
, ype 2
diabe es
(5–7)
, he me abolic synd ome
(8)
,inflamma ion
(9–11)
and cogni i e decline
(12,13)
, and hey may also con ibu e o
heal hy ageing
(14)
and weigh main enance
(15)
. Thus, he scien-
ific e idence o heal h benefi s is encou aging, and he No dic
Nu i ion Recommenda ions
(16)
ad ise o inc ease consump-
ion o be ies as pa o a balanced heal hy die .
Due o low suga :acid a io and high con en o polyphenols,
sou and bi e as e is ypical o many be ies. Suga , such as
suc ose o high- uc ose co n sy up, is commonly used o
inc ease accep abili y o be y p oduc s. Al hough suga helps
o inc ease be y consump ion, i may comp omise he nu i-
ional and heal h benefi s o be ies, e.g. by inducing pos p an-
dial hype glycaemia. The e o e, i is impo an o unde s and he
me abolic consequences o suga consumed wi h be ies.
Ou p e ious s udies ha e shown ha be ies may imp o e
pos p andial glycaemic con ol in heal hy subjec s when
Abb e ia ions: i.d., in e nal diame e ; SGLT1, sodium glucose co- anspo e 1.
*Co esponding au ho : R. Tö önen, email ii a. o onen@ue .fi
© The Au ho (s) 2017. This is an Open Access a icle, dis ibu ed unde he e ms o he C ea i e Commons A ibu ion licence (h p://c ea i e-
commons.o g/licenses/by/4.0/), which pe mi s un es ic ed e-use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal wo k is
p ope ly ci ed.
JNS
JOURNAL OF NUTRITIONAL SCIENCE
1
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consumed wi h suc ose. The pos -meal fluc ua ion o blood
glucose and insulin concen a ions induced by suc ose was alle-
ia ed when suc ose was consumed wi h a mix u e o ou be -
ies (bilbe ies, blackcu an s, c anbe ies and s awbe ies)
(17,18)
.
In addi ion, blackcu an s and lingonbe ies, ei he as whole
be ies o nec a s, imp o ed he pos p andial glycaemic and
insulinaemic esponses o suc ose
(19)
. These esponses a e con-
sis en wi h delayed diges ion o suc ose and/o slowe abso p-
ion o glucose caused by be y polyphenols.
Suc ose is a disaccha ide composed o glucose and uc ose.
In he p esence o low pH and hea , suc ose is in e ed, i.e. he
glycosidic linkage is hyd olysed in an i e e sible eac ion,
eleasing glucose and uc ose. In ou p e ious s udies
(17–19)
,
suc ose was added o he be y p oduc s jus be o e he p od-
uc was consumed, o a oid in e sion. Howe e , du ing p o-
cessing and s o age o be y p oduc s wi h added suc ose,
in e sion o suc ose is likely o occu , due o he low pH o
be ies. The e o e, unde s anding he e ec s o be ies on
he glycaemic esponse induced by in e ed suc ose is o p ac-
ical impo ance. The pu pose o his s udy was o in es iga e
he pos p andial glycaemic and insulinaemic esponses o
blackcu an (Ribes nig um) and lingonbe y (Vaccinium
i is-idaea) nec a s con aining in e ed suc ose, added as glu-
cose and uc ose, in heal hy subjec s.
Me hods
Pa icipan s
Volun ee s we e ec ui ed by an announcemen in a local
newspape in he a ea o Kuopio, Finland. The inclusion c i-
e ia we e as ollows: heal hy men o women, age 25–69
yea s, BMI 20–28 kg/m
2
, no diabe es o o he ch onic disease,
no smoking, no an ibio ic medica ion wi hin he pas 3
mon hs, and no blood dona ion wi hin he pas mon h. A
he sc eening isi , hei heal h s a us and medical his o y
was assessed by using a s uc u ed in e iew, and by an h opo-
me ic and labo a o y measu emen s ( ou ine haema ological
measu emen s, as ing plasma glucose, lipids, hy oid-
s imula ing ho mone, alanine amino ans e ase and c ea inine).
The s udy was conduc ed acco ding o he guidelines laid
down in he Decla a ion o Helsinki and all p ocedu es in ol -
ing human subjec s we e app o ed by he Resea ch E hics
Commi ee o he Hospi al Dis ic o No he n Sa o
(Finland). W i en in o med consen was ob ained om all
pa icipan s. The s udy was egis e ed a clinical ials.go as
NCT02743130.
Tes p oduc s
Blackcu an and lingonbe y nec a s we e p epa ed by
Eckes-G anini Finland Oy Ab om indus ial juice concen-
a es, wi h he final juice concen a ion o 50 %. Sodium
benzoa e was added as a p ese a i e, and he nec a s we e
pas eu ised and s o ed a +5°C. Suga composi ion was ana-
lysed by HPLC. In he esea ch labo a o y, glucose (Glucosum
anhyd icum Ph.Eu .; Vi abalans Oy) and uc ose (F uisana;,
Danisco Swee ene s Oy) we e added o achie e final
concen a ions o 17·5 g in 300 ml nec a (Table 1).
Acco ding o Eu opean legisla ion, p oduc s ob ained by add-
ing wa e and suga s o concen a ed ui juice a e defined as
nec a s
(20)
. A e e ence d ink con ained 17·5 g glucose and
17·5 g uc ose in 300 ml ap wa e . Polyphenols (an hocya-
nins, p oan hocyanidins, fla onols and phenolic acids) in he
nec a s we e analysed be o e addi ion o he suga s.
Re e ence s anda ds o an hocyanins (delphinidin-3-
glucoside, delphinidin-3- u inoside, cyanidin-3-galac oside,
cyanidin-3-glucoside, cyanidin-3-a abinoside, cyanidin-3-
u inoside, pe unidin-3-glucoside, peonidin-3-glucoside,
mal idin-3-glucoside) we e pu chased om Ex asyn hese.
T iplica e samples we e weighed and dilu ed wi h a mix u e
o me hanol, ace ic acid and wa e (65:4:31) p io o HPLC
analysis. An hocyanins we e sepa a ed on a 150 mm, 4·6
mm in e nal diame e (i.d.), 5 µm, Gemini C18 column wi h
a C18 gua d column using a g adien elu ion o ace oni ile
in o 5 % o mic acid acco ding o Hells öm e al.
(21)
.
De ec ion wa eleng h was se a 518 nm and UV spec um
was eco ded be ween 190 and 600 nm. Iden ifica ion was
based on e e ence s anda ds, UV spec a and li e a u e.
Au hen ic s anda ds we e used o quan ifica ion when a ail-
able; i no , he co esponding an hocyanidin-3-glucoside was
used (e.g. pe unidin-3- u inoside was quan ified as
pe unidin-3-glucoside).
Re e ence s anda ds o p oan hocyanidins (ca echin, epica-
echin and p ocyanidin B2) we e pu chased om
Ex asyn hese. P ocyanidin oligome s (deg ee o polyme isa-
ion 3–10) we e om Plan a Analy ica; he pu i y was de e -
mined wi h hiolysis
(22)
. T iplica e samples (10 ml) we e
pu ified wi h polyamide Supelco Disco e y DPA-6S
(Sigma-Ald ich Chemie Inc.)
(22)
. HPLC analyses we e ca ied
ou wi h an Agilen 1100 se ial de ice using a Phenomenex
Luna HILIC column (250 mm, 4·6 mm i.d., 5 µm) wi h diol
phase. The elu ion sol en s we e ace oni ile + ace ic acid =
98 + 2 (A), me hanol + ace ic acid + wa e = 95 + 2 + 3 (B)
and dime hyl o mamide + ace ic acid + wa e = 85 + 2 + 13
(C). P oan hocyanidins we e sepa a ed a he flow a e o 1
ml/min acco ding o he ollowing e na y g adien : 95 % A
and 5 % B as ini ial condi ions; changing o 40 % A and 60
% B in 45 min; isoc a ic elu ion (40 % A and 60 % B) o
5 min; changing o 60 % B and 40 % C in 1 min; changing
o 20 % B and 80 % C in 2 min; isoc a ic elu ion (20 % B
and 80 % C) o 7 min; changing o 80 % A and 20 % B in
4 min; changing o 95 % A and 5 % B in 1 min. Equalising
ime be ween he injec ions was 10 min. Fluo escence de ec-
ion wi h exci a ion wa eleng h o 280 nm and emission wa e-
leng h 323 nm was used o he de e mina ion.
Table 1. Suga composi ion o he es p oduc s
Re e ence
d ink
Blackcu an
nec a
Lingonbe y
nec a
Glucose, na u al (g) 4·68 4·50
Glucose, added (g) 17·50 12·82 13·00
F uc ose, na u al (g) 5·07 3·81
F uc ose, added (g) 17·50 12·43 13·69
Juice (ml) 300 300
Wa e (ml) 300
2
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Fla onol glycosides we e hyd olysed by efluxing he sam-
ples in 1·2M-HCl in 50 % aqueous me hanol o 1 h acco ding
o He og e al.
(23)
. Fla onol aglycones we e iden ified and
quan ified using an Agilen 1100 Se ies HPLC equipped
wi h a diode a ay de ec o . The analy ical column was a
No a Pak C18 (150 mm, 3·9 mm i.d., 4 µm, Wa e s) p o ec ed
wi h he manu ac u e ’s p ecolumn. The mobile phase con-
sis ed o 0·05 M-phospha e bu e (A) a pH 2·4 and me hanol
(B) (5–60 % B in 50 min ollowed by 60–90 % B in 6 min).
Iso hamne in, kaemp e ol, my ice in and que ce in we e quan-
ified a 370 nm. Fo iden ifica ion pu poses, UV/ isable
spec a we e eco ded a 190–600 nm.
Re e ence s anda ds o phenolic acids (ca eic acid, e ulic
acid, p-couma ic acid, cinnamic acid) we e pu chased om
Sigma-Ald ich. T iplica e samples we e fil a ed and analysed
by Agilen 1290 Infini e UHPLC equipped wi h a diode
a ay de ec o . Phenolic acids we e sepa a ed a a flow a e
o 0·4 ml/min on a 50 mm, 2·1 mm i.d., 1·8 µm, Zo bax
Eclipse Plus C18 column using a g adien elu ion o ace o-
ni ile (B) in o 50 mM-phospho ic acid (pH 2·5, A): 95 % A
and 5 % B as ini ial condi ions; isoc a ic elu ion o 1·2 min;
changing o 85 % A and 15 % B in 4·05 min; changing o
80 % A and 20 % B in 5·75 min; changing o 50 % A and
50 % B in 5 min; isoc a ic elu ion o 1·2 min; changing o
95 % A and 5 % B in 0·8 min. De ec ion wa eleng hs we e
280 nm and 329 nm, and UV spec um was eco ded a
190–400 nm. Phenolic acids we e iden ified acco ding o
hei UV spec a and quan ified as he co esponding aglyconic
o ms.
S udy design
The andomised, con olled, c osso e s udy was ca ied ou
single-blinded o he labo a o y pe sonnel and analyses.
Each subjec pa icipa ed in h ee 2 h es s on sepa a e isi s,
a leas 3 d apa . The es p oduc s we e se ed in a ando-
mised o de .
The pa icipan s we e ins uc ed o keep hei die , body
weigh and li ing habi s cons an du ing he s udy, abs ain
om alcohol o 2 d be o e he es , and o e ain om in en-
si e physical ac i i y o 12 h be o e he es . The day be o e
he s udy isi , hey we e ad ised no o consume any be ies
o be y p oduc s. In he e ening be o e he isi , hey we e
ins uc ed o consume a meal o choice and epea ha meal
be o e e e y isi . Body weigh was measu ed a e e y isi .
The es s we e ca ied ou in he mo ning a e a 10–12 h
o e nigh as . Fo collec ion o enous blood samples, an in a-
enous ca he e (BD Venflon™P o 18GA, 1·3 × 32 mm;
Beck on Dickinson) was inse ed in an an ecubi al ein o
he o ea m. Baseline blood samples we e d awn a as ing
(0 min) and o he samples a 15, 30, 45, 60, 90 and 120 min
a e s a ing o consume he es p oduc . The samples o
plasma glucose measu emen s we e collec ed in ci a e–fluo ide
ubes and o plasma insulin measu emen s in EDTA ubes
kep on ice. Plasma was immedia ely sepa a ed by cen i uga ion
a +4°C and s o ed a −70°C.
Glucose concen a ions we e analysed wi h he hexokinase
me hod using Konelab Sys em eagen s and a Konelab 20
XTi clinical chemis y analyse (The mo Fishe Scien ific).
The in a- and in e -assay CV we e 2·7 and 1·8 %, espec -
i ely. Insulin was measu ed wi h a chemiluminome ic immuno-
assay using Liaison® Insulin eagen s (DiaSo in Inc.) and a
DiaSo in Liaison® analyse (DiaSo in Deu chland GmbH).
The in a- and in e -assay CV we e 3·7and3·8 %, espec i ely.
Calcula ions and s a is ical analyses
The da a we e analysed by using IBM SPSS S a is ics o
Windows, e sion 21.0 (IMB Co p.), and a e exp essed as
means and s anda d de ia ions o s anda d e o s, as indi-
ca ed. Linea mixed-e ec s modelling was used o compa e
he e ec s o he es p oduc s. His og ams we e used o
checking he no mali y o model esiduals. Values o P<
0·05 we e conside ed o be s a is ically significan .
The s a is ical significance o he p oduc × ime in e ac ion
was es ed in he mixed-model analysis using pa icipan as a
andom ac o and p oduc × ime and hei main e ec s as
fixed ac o s. When he p oduc × ime in e ac ion was s a is-
ically significan , he di e ences be ween he e e ence d ink
and be y nec a s a indi idual ime poin s we e es ed by
pos hoc analysis wi h Sidak adjus men o mul iple
compa isons.
The maximum inc eases om he baseline concen a ions
we e calcula ed by sub ac ing he as ing alue om he high-
es alue. G aphPad P ism 5.03 o Windows so wa e
(G aphPad So wa e, Inc.) was used o calcula ion o he gly-
caemic p ofile o each subjec and es p oduc acco ding o
Rosén e al.
(24)
, by di iding he ime (min) du ing which he
plasma glucose was abo e he as ing concen a ion wi h he
inc emen al peak alue (mmol/l). The inc emen al AUC
we e calcula ed by using G aphPad P ism so wa e, igno ing
he a ea below he baseline (0 min) concen a ion. The di e -
ences be ween he e e ence and be y nec a s we e es ed
using pa icipan as a andom ac o and p oduc as a fixed
ac o in he mixed-model analysis.
Resul s
A e sc eening wen y-six olun ee s, hi een emale and fi e
male pa icipan s we e en olled o he s udy, and hey all com-
ple ed he h ee isi s. The a e age ime be ween isi s was 7·6
( ange 3–17) d. The basic cha ac e is ics o he pa icipan s a e
p esen ed in Table 2. The body weigh s emained s able
Table 2. Basic cha ac e is ics o he pa icipan s (n13 emale and n5
male)
(Mean alues, s anda d de ia ions and anges)
Mean SD Range
Age (yea s) 59 7 40–68
Weigh (kg) 66·99·453–86
BMI (kg/m
2
)23·92·220–28
Fas ing plasma glucose (mmol/l) 5·40·34·9–6·1
Fas ing plasma o al choles e ol (mmol/l) 5·30·83·9–7·1
Fas ing plasma LDL-choles e ol (mmoll/l) 3·00·91·7–5·7
Fas ing plasma HDL-choles e ol (mmol/l) 2·10·41·2–2·8
Fas ing plasma TAG (mmol/l) 0·90·30·5–1·6
3
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h oughou he s udy pe iod. The mean body weigh s we e
67·5(
SD 9·6), 67·6(SD 9·5) and 67·3(SD 9·3) kg o he e e -
ence, blackcu an nec a and lingonbe y nec a isi s,
espec i ely.
The e we e no di e ences in he imes spen d inking he
es p oduc s. The e e ence d ink was inges ed in 2:36 (SD
0:59), blackcu an nec a in 2:37 (SD 1:04) and lingonbe y
nec a in 2:31 (SD 0:59) min:s.
Glucose esponse
Since one pa icipan had an abno mal glucose esponse cu e,
he glucose a iables ha e been calcula ed o se en een pa i-
cipan s. The pos p andial glucose esponses a e p esen ed in
Fig. 1. The p oduc × ime in e ac ion was s a is ically signifi-
can (P=0·003). In compa ison wi h he e e ence, inges ion
o he blackcu an nec a esul ed in lowe glucose concen a-
ions a 15 and 30 min and a highe concen a ion a 60 min.
A e he blackcu an nec a , he maximum inc ease om
baseline was a enua ed by 33 %, and he glycaemic p ofile
was imp o ed by 84 % (Table 3). The lingonbe y nec a
had no s a is ically significan e ec on he glucose a iables.
Insulin esponse
The pos p andial insulin esponses a e p esen ed in Fig. 2. The
p oduc × ime in e ac ion was highly significan (P<0·001).
In compa ison wi h he e e ence, lowe insulin concen a ions
we e obse ed a 15 min o bo h nec a s and a 30 min o
he lingonbe y nec a . A 60 min, he insulin concen a ion
emained highe a e he blackcu an nec a . In compa ison
wi h he e e ence, he maximum inc ease om he baseline
was a enua ed by 21 % a e he lingonbe y nec a
(Table 3). A e he blackcu an nec a , he maximum inc ease
om he baseline was 13 % smalle han a e he e e ence,
bu he di e ence is no s a is ically significan .
Polyphenol composi ion
An hocyanins and p oan hocyanidins we e he majo polyphe-
nols in he nec a s, and fla onols (que ce in) and phenolic
acids (p-couma ic, ca eic and e ulic acids) we e de ec ed in
smalle quan i ies (Table 4).
The con en o an hocyanins was much highe in he black-
cu an nec a (57·9 mg/100 g) han in he lingonbe y nec a
(4·5 mg/100 g) (Table 4). The an hocyanin p ofile o he
blackcu an nec a was domina ed by u inosides o delphini-
din and cyanidin ollowed by glucosides o he same an hocya-
nidins. Ru inosides o pe unidin and peonidin we e iden ified
as mino an hocyanins. In he lingonbe y nec a , cyanidin-3-
galac oside was he mos abundan , and cyanidin-3-a abinoside
and cyanidin-3-glucoside we e p esen in smalle amoun s. In
addi ion, aces o delphinidin and mal idin de i a i es we e
de ec ed, p obably o igina ing om con amina ion o he
lingonbe y juice concen a e wi h bilbe y.
The lingonbe y nec a con ained mo e p oan hocyanidins
(71·5 mg/100 g) han he blackcu an nec a (37·1 mg/100
g) (Table 4). Oligome ic p oan hocyanidins (deg ee o poly-
me isa ion 2–10) domina ed in he lingonbe y nec a , and
35 % o he dime s we e A- ype (could no be es ima ed o
o he oligome s). On he o he hand, 90 % o he
Fig. 1. Plasma glucose concen a ions a e inges ion o he es p oduc s: e -
e ence (▲); blackcu an nec a (●); lingonbe y nec a (○). Values a e means
(n17), wi h s anda d e o s ep esen ed by e ical ba s. P=0·003 o p od-
uc × ime in e ac ion in he mixed-model analysis. Mean alue o he black-
cu an nec a was signi ican ly di e en om ha o he e e ence a an
indi idual ime poin : * P<0·05, ** P<0·01 ( pos hoc analysis wi h Sidak
adjus men ).
Table 3. Glucose and insulin a iables a e consump ion o he es p oduc s
(Mean alues and s anda d de ia ions)
Re e ence Blackcu an nec a Lingonbe y nec a
Mean SD Mean SD Mean SD Mixed model analysis: P
Glucose†
Fas ing (mmol/l) 5·30·35·30·35·10·4>0·05
Maximum inc ease (mmol/l) 2·41·01·6** 0·92·20·50·007
AUC (mmol/l × min) 73·337·755·234·573·929·50·055
Glycaemic p o ile (min/(mmol/l)) 25·27·246·4*** 22·829·76·4<0·001
Insulin‡
Fas ing (mU/l) 5·02·05·73·45·42·8>0·05
Maximum inc ease (mU/l) 27·912·624·314·221·9* 12·10·040
AUC (mU/l × min) 1089 492 1117 482 958 436 0·090
Mean alue was signi ican ly di e en om ha o he e e ence: * P<0·05, ** P<0·01, *** P<0·001 (pos hoc analysis wi h Sidak adjus men ).
†n17.
‡n18. To con e insulin in mU/l o pmol/l, mul iply by 6·945.
4
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p oan hocyanidins in he blackcu an nec a we e highly poly-
me ised (deg ee o polyme isa ion >10).
Discussion
Suc ose is commonly used o swee ening be y juices and
o he be y p oduc s. When he me abolic e ec s o
suc ose-swee ened be y p oduc s a e e alua ed, in e sion o
suc ose should be conside ed. In his s udy we in es iga ed
he e ec s o be y nec a s swee ened wi h glucose and uc-
ose, ep esen ing comple ely in e ed suc ose, on pos p andial
glucose me abolism. Ou esul s show ha he ea ly pos p an-
dial glycaemic esponses induced by in e ed suc ose in he
be y nec a s a e simila o hose obse ed in ou p e ious
s udies
(18,19)
whe e in e sion o suc ose was a oided.
The e a e e y ew s udies compa ing he glycaemic e ec o
suc ose and in e ed suc ose. The s udy o Ellwood e al.
(25)
ound no significan di e ence in he glycaemic esponse
be ween a load o suc ose o in e ed suc ose. Compa able
glycaemic esponses ha e also been epo ed o suc ose and
high- uc ose co n sy up, a swee ene consis ing o ee uc-
ose and glucose
(26–29)
. Howe e , i is possible ha suc ose
gi en in he o m o a be e age in hese s udies was hyd o-
lysed, as epo ed by Le e al.
(29)
. The p esen s udy shows
ha in e ed suc ose induces pos p andial glucose and insulin
esponses compa able wi h hose o in ac suc ose. The
esponse cu es o he e e ence d ink con aining 17·5 g glu-
cose and 17·5 g uc ose a e p ac ically simila o hose
epo ed p e iously o 35 g suc ose
(18,19)
. The maximum
inc eases om he baseline we e 2·4 and 2·3 mmol/l o glu-
cose and 27·9 and 33·4 mU/l o insulin in his s udy and in
he p e ious s udy
(19)
, espec i ely. The alues o he gly-
caemic p ofile in hese s udies we e 25·2 and 29·6 min/
(mmol/l), espec i ely.
Blackcu an in he o m o a nec a (50 % juice) signifi-
can ly a enua ed he in e ed suc ose-induced pos meal ise
o plasma glucose concen a ion and imp o ed he glycaemic
p ofile, whe eas he e ec on insulin esponse was less p o-
nounced. The lowe glycaemic esponse was e iden du ing
he ea ly pos p andial phase, and in he la e phase he declines
o he glucose and insulin concen a ions we e less s eep han
a e he e e ence. These esul s a e iden ical o hose de ec ed
p e iously o suc ose added o a simila blackcu an nec-
a
(19)
. The maximum inc ease o glucose concen a ion
om he baseline (1·6 mmol/l o bo h) and he alues o
he glycaemic p ofile (46·4 and 46·1 min/(mmol/l)) we e simi-
la in his s udy and in he p e ious s udy.
The glycaemic esponse induced by in e ed suc ose in he
lingonbe y nec a did no di e om he e e ence. Also in
he p e ious s udy
(19)
, suc ose in he lingonbe y nec a had
no e ec on he ea ly glucose esponse. Howe e , in ac
suc ose in he lingonbe y nec a p oduced a mo e sus ained
la e -phase glucose esponse, p e en ed he suc ose-induced
la e pos p andial hypoglycaemia and he eby imp o ed he gly-
caemic p ofile
(19)
. These e ec s we e no obse ed in he p e-
sen s udy. Fu he s udies a e needed o e i y and explain his
di e ence. In e ed suc ose in he lingonbe y nec a signifi-
can ly dec eased he ea ly insulin esponse. A enua ion o
pos p andial insulin esponse by lingonbe ies, wi h li le o
no e ec on glucose esponse, was p e iously obse ed
when whole-be y pu ée was consumed wi h suc ose o
whi e whea b ead
(19,30)
.
We ha e p e iously sugges ed ha be ies may modula e he
pos p andial glycaemic esponse o suc ose by educing diges-
ion o suc ose and/o abso p ion o glucose, and ha
Fig. 2. Plasma insulin concen a ions a e inges ion o he es p oduc s: e -
e ence (▲); blackcu an nec a (●); lingonbe y nec a (○). Values a e means
(n18), wi h s anda d e o s ep esen ed by e ical ba s. P<0·001 o p oduc
× ime in e ac ion in he mixed-model analysis. Mean alue o he blackcu -
an nec a was signi ican ly di e en om ha o he e e ence a an indi idual
ime poin : ** P<0·01, *** P<0·001 (pos hoc analysis wi h Sidak adjus men ).
Mean alue o he lingonbe y nec a was signi ican ly di e en om ha o he
e e ence a an indi idual ime poin : †P<0·05, †† P<0·01 (pos hoc analysis
wi h Sidak adjus men ). To con e insulin in mU/l o pmol/l, mul iply by 6·945.
Table 4. Polyphenol con en s in he nec a s (mg/100 g)
(Mean alues and s anda d de ia ions o iplica e analyses)
Blackcu an
nec a
Lingonbe y
nec a
Mean SD Mean SD
An hocyanins, o al con en 57·90·84·50·2
Delphinidin-3-glucoside 5·93 0·09 0·18 0·02
Delphinidin-3- u inoside 28·50·40 –
Delphinidin de i a i es (as D-3-Glc) –0·84 0·05
Cyanidin-3-galac oside –2·19 0·09
Cyanidin-3-glucoside 2·63 0·04 0·45 0·02
Cyanidin-3-a abinoside –0·65 0·03
Cyanidin-3- u inoside 19·50·30 –
Pe unidin-3- u inoside 1·01 0·04 –
Peonidin-3- u inoside 0·33 0·03 –
Mal idin-3-glucoside –0·17 0·01
P oan hocyanidins, o al con en 37·10·671·53·0
P1 0·50 0·014 3·43 0·19
P2 0·67 0·021 9·49 0·24
P3 0·49 0·039 7·53 0·22
P4–P6 1·26 0·13 15·20·6
P7–P10 0·72 0·061 6·78 0·65
>P10 33·50·529·11·4
Fla onols, o al con en 1·70·14·00·1
Que ce in 1·70·14·00·1
Phenolic acids, o al con en 1·72 0·02 1·17 0·01
Ca eic acid 0·741 0·011 0·24 0·001
Fe ulic acid 0·22 0·002 0·17 0·001
p-Couma ic acid 0·76 0·010 0·37 0·006
Cinnamic acid –0·39 0·002
P, deg ee o polyme isa ion.
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polyphenols may be he ac i e compounds
(17–19)
. In he small
in es ine, suc ose is hyd olysed o glucose and uc ose by
α-glucosidase, which is known o be inhibi ed by be y ex ac s
and polyphenols in i o
(31–33)
. In he p esen s udy, howe e ,
α-glucosidase ac i i y plays no ole, because suga was inges ed
as ee glucose and uc ose. The e o e, he lowe glycaemic
esponse obse ed a e consump ion o he blackcu an nec-
a canno be explained by inhibi ion o α-glucosidase ac i i y.
Inhibi ion o glucose abso p ion om he small in es ine is
ano he po en ial mechanism o glycaemic con ol a e a meal.
The in es inal abso p ion o glucose is mainly media ed by wo
anspo e s, he sodium glucose co- anspo e 1 (SGLT1) in
he apical memb ane and GLUT2 in he basola e al memb ane
o en e ocy es
(34,35)
. Polyphenols and polyphenol- ich ex ac s
a e known o in e ac wi h hese anspo e s and may he eby
egula e he a e o glucose abso p ion
(36,37)
.
In acco dance wi h p e ious s udies
(38–41)
, he polyphenol
p ofile o he blackcu an nec a was cha ac e ised by he
abundance o an hocyanins, especially by u inosides and glu-
cosides o delphinidin and cyanidin, and highly polyme ised
p oan hocyanidins, while lingonbe y nec a was cha ac e ised
by oligome ic and polyme ic p oan hocyanidins, wi h lowe
con en s o an hocyanins and fla onols. Ou p e ious
(19)
and p esen esul s show ha an hocyanin- ich blackcu an
is mo e po en han lingonbe y in lowe ing pos -meal gly-
caemia p oduced by in ac o in e ed suc ose, espec i ely.
The e a e no p e ious s udies on he in e ac ion o blackcu -
an o i s componen s wi h glucose abso p ion. Howe e , in
i o s udies ha e demons a ed ha o he an hocyanin- ich be -
ies a e able o influence in es inal glucose anspo . In human
in es inal Caco-2 cells, s awbe y ex ac inhibi ed glucose up ake
om heapicalsidein o hecellsand heGLUT2- acili a ede flux
on he basola e al side
(42)
. Pela gonidin-3-glucoside, he main
an hocyanin in s awbe y, was esponsible o 26 % o he o al
inhibi ion. Mo eo e , an hocyanin- ich be y ex ac dec eased
bo h Na-dependen and Na-independen glucose up ake and
educed SGLT1 mRNA and GLUT2 mRNA exp ession in
Caco-2 cells
(43)
. In e es ingly, cyanidin-3-glucoside and
cyanidin-3- u inoside we e ound o inhibi bo h ypes o glucose
anspo . These compounds and delphinidin glycosides we e he
main an hocyanins o he blackcu an nec a o ou s udy. A
maquibe yex ac ichindelphinidinglycosidesando he an ho-
cyaninsdec easedpos p andialglucoseandinsulinconcen a ions
a e s a ch consump ion, and SGLT1 inhibi ion by delphinidin
was sugges ed o be he p ima y mechanism
(44)
.
In conclusion, he pos p andial glycaemic and insulinaemic
esponses induced by in e ed suc ose a e simila o hose
obse ed p e iously o in ac suc ose. Modula ion o he
ea ly glycaemic esponse by be y nec a s is p ac ically simila
o in ac and in e ed suc ose, sugges ing ha in e sion has
no majo impac on glycaemic esponse o suc ose-swee ened
be y p oduc s. The a enua ed glycaemic esponse o he
blackcu an nec a con aining in e ed suc ose may be
explained by inhibi ion o in es inal abso p ion o glucose by
blackcu an an hocyanins. In u u e s udies, he e ec s o
o he an hocyanin- ich be ies, such as bluebe ies, bilbe ies
and chokebe ies, on suc ose-induced pos p andial glycaemia
should be in es iga ed.
Acknowledgemen s
The au ho s would like o hank he s udy pa icipan s o hei
ime and e o , Hannele Heinonen and Ee a Lajunen o hei
skil ul echnical assis ance in he clinical s udy, and
Eckes-G anini Finland Oy Ab o p o iding he nec a s.
R. T. and K. K. designed he esea ch, R. T. conduc ed he
esea ch, analysed he da a, w o e he pape and had p ima y
esponsibili y o he final con en . J. H. and P. M. ca ied
ou he polyphenol analyses and con ibu ed o w i ing he
pape . All au ho s ead and app o ed he final manusc ip .
The p esen s udy was suppo ed by unding om
Finnsuga L d. K. K. is an employee o Finnsuga
L d. R. T., J. H. and P. M. decla e no conflic o in e es .
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