Biochemical and Molecular Actions of Nutrients

Tea Catechins with a Galloyl Moiety Suppress Postprandial

Hypertriacylglycerolemia by Delaying Lymphatic Transport
of Dietary Fat in Rats
Ikuo Ikeda,1 Koichi Tsuda, Yuko Suzuki,* Makoto Kobayashi,* Tomonori Unno,*
Hiroko Tomoyori, Hitomi Goto, Yayoi Kawata, Katsumi Imaizumi, Ayumu Nozawa,*
and Takami Kakuda*
Laboratory of Nutrition Chemistry, Department of Bioscience and Biotechnology, Faculty of Agriculture,
Graduate School of Kyushu University, Fukuoka 812-8581, Japan; *Central Research Institute, ITO EN, Ltd.,
Shizuoka 421-0516, Japan; and Department of Environmental and Symbiotic Sciences, Faculty of D
o
Environmental and Symbiotic Sciences, Prefectural University of Kumamoto, Kumamoto 862-8502, Japan w
nl
oa
ABSTRACT Tea catechins, ( )-epicatechin (EC), ( )-epigallocatechin (EGC), ( )-epicatechin gallate (ECG), and de
( )-epigallocatechin gallate (EGCG), have been shown to be epimerized to ( )-catechin (C), ( )-gallocatechin d
(GC), ( )-catechin gallate (CG), and ( )-gallocatechin gallate (GCG), respectively, during heat treatment. In this fro
study, we examined the effect of tea catechins rich in ECG and EGCG and heat-treated tea catechins rich in CG m
and GCG on postprandial hypertriacylglycerolemia in rats. Both tea catechins and heat-treated tea catechins jn.
14 nu
suppressed postprandial hypertriacylglycerolemia. Lymphatic recovery of C-trioleoylglycerol in rats cannulated in trit
the thoracic duct was delayed by the administration of tea catechins and heat-treated tea catechins. Tea io
catechins and heat-treated tea catechins had the same effect on all variables tested. These catechin n.
preparations dose- dependently inhibited the activity of pancreatic lipase in vitro. When purified catechins were or
used, only those with a galloyl moiety inhibited the activity of pancreatic lipase. These results suggest that g
at
catechins with a galloyl moiety suppress postprandial hypertriacylglycerolemia by slowing down triacylglycerol
Ku
absorption through the inhibition of pancreatic lipase. Because postprandial hypertriacylglycerolemia is a risk n
factor for coronary heart disease, our results suggest that catechins with a galloyl moiety may prevent this mi
disease. J. Nutr. 135: 155159, 2005. ng
In
KEY WORDS: catechins postprandial hypertriacylglycerolemia pancreatic lipase rats tea sti
tut
Tea catechins consist mainly of 4 derivatives; ( )-epicat- tea extract slows or inhibits fat absorption in the intestine and e
2
echin (EC), ( )-epigallocatechin (EGC), ( )-epicatechin suppresses postprandial hypertriacylglycerolemia, the effect of
gallate (ECG), and ( )-epigallocatechin gallate (EGCG) has never been experimentally tested. Tea catechins are Bo
(Fig. 1). These catechins, which are contained in green tea, thought to be responsible for the lipase inhibitor in green tea ta
oolong tea, and black tea, have been shown to have hypocho- extract. However, Han et al. (18) reported that tea saponins, ny,
lesterolemic, antiatherogenic, antiobesity, antioxidative, and not catechins, inhibited pancreatic lipase activity in vitro. In C
anticarcinogenic activities (110). We previously showed that this study, the effect of tea catechins on postprandial hyper- A S
tea catechins inhibit intestinal absorption of cholesterol in rats triacylglycerolemia was examined in rats, and the underlying on
(11,12). It also has been reported that tea catechins prevent mechanism was clarified in vitro and in vivo. We used rats as Fe
LDL oxidation (1315) and suppress atheroma formation in a model of postprandial hypertriacylglycerolemia, because the br
apoE knockout mice (5). These observations suggest that tea processes of digestion and absorption of dietary ua
catechins are effective in preventing coronary heart disease. triacylglycerols are similar in rats and humans.
Because postprandial hypertriacylglycerolemia is a risk factor Canned and bottled tea beverages are widely consumed in
for coronary heart disease (16), its suppression by food com- Asian countries. For example, 1.01 109 L canned and
ponents may effectively prevent the disease. Juhel et al. (17) bottled tea beverages were produced in 2000 in Japan. About
showed that a green tea extract inhibited gastric and pancre- 50% of tea catechins in these beverages are epimerized by heat
atic lipases in vitro. Although the results suggest that a green treatment to ( )-catechin (C), ( )-gallocatechin (GC), ( )-
catechin gallate (CG), and ( )-gallocatechin gallate (GCG)
(Fig. 1) (19,20). Although many studies have been conducted
1
To whom correspondence should be addressed. on physiological functions of tea catechins, studies on heat-
E-mail: iikeda@agr.kyushu-u.ac.jp.
2
Abbreviations used: C, ( )-catechin; CG, ( )-catechin gallate; EC, ( )-
treated tea catechins are scarce. In this study, the effect of
epicatechin; ECG, ( )-epicatechin gallate; EGC, ( )-epigallocatechin; EGCG, heat-treated tea catechins on postprandial hypertriacylglycer-
( )-epigallocatechin gallate; GC, ( )-gallocatechin; GCG, ( )-gallocatechin gal- olemia was also compared with tea catechins.
late; TES, N-tris(hydroxymethyl)methyl-2-aminoethane sulfonic acid.

0022-3166/05 $8.00 2005 American Society for Nutritional Sciences.

Manuscript received 4 July 2004. Initial review completed 22 July 2004. Revision accepted 21 October 2004.

155
156
IKEDA ET AL.

FIGURE 1 Chemical structures of tea catechins and their corre-
sponding epimers.
MATERIALS AND METHODS
Purified catechins, EC, EGC, ECG, EGCG, C, GC, CG, and
GCG were obtained from Wako Pure Chemicals. Purities of all of
these catechins were 98%. THEA-FLAN 90S, a decaffeinated
green tea powder provided from ITO EN, was used as a mixture of
tea catechins. This product was rich in EGCG and ECG. Heat-
treated tea catechins were prepared by autoclaving the catechin
mixture at
120C for 5 min. The composition of catechins in tea catechins and
heat-treated tea catechins is shown in Table 1. Heat-treated tea
catechins prepared from tea catechins contained increased amounts
14
of GCG and CG. Tri[1- C]oleoylglycerol (2.0 GBq/mmol) was
obtained from Amersham Pharmacia Biotech.
Postprandial hypertriacylglycerolemia in rats. Male Wistar rats
(8 wk old, SPF, Japan SLC) were fed a commercial nonpurified diet
(CE-2; CLEA Japan) for 1 wk and deprived of food overnight. After
blood was withdrawn from the jugular vein as 0 time, a solution
containing tea catechins or heat-treated tea catechins [100 mg/(5 mL
deionized water kg body weight)] was given orally. Deionized water
was given in the control group. Immediately after administration, a
lipid emulsion containing 200 g/L soybean oil, 12 g/L egg lecithin,
and 22.5 g/L glycerin was given orally, at a dose of 10 mL/kg (21).
Blood (100 L) was withdrawn at 1, 2, 3, 4, 6, and 8 h after
administration via the jugular vein. After the separation of serum,
triacylglycerol was measured with a commercial kit (Triglyceride E
test Wako; Wako Pure Chemicals).
14
Lymphatic recovery of C-trioleoylglycerol in rats cannulated in
the thoracic duct. Nine-wk-old male Sprague Dawley rats cannu-
lated in the thoracic duct were given i.g. 3 mL of a test emulsion
14
containing C-trioleoylglycerol with or without catechins. After
administration, lymph was collected for 24 h. The test emulsion (3
mL) contained 200 mg sodium taurocholate (Nacalai tesque), 50 mg
fatty-acid-free bovine serum albumin 14fraction V (Bayer), 200 mg
trioleoylglycerol (Sigma), and 37 kBq C-trioleoylglycerol in deion-
ized water. When tea catechins and heat-treated tea catechins were
administered, these catechins were added to the emulsion at 100 mg
in 3 mL test emulsion, respectively. Operations and maintenance of
rats and all other procedures were performed as described previously
(22,23).
All rat studies were carried out under the guidelines for animal
experiments of the Faculty of Agriculture, Graduate School Kyushu
University and Central Research Institute, ITO EN, and Law 105 and
Notification 6 of the government of Japan.
Micellar solubility of hydrolysis products of triacylglycerol in
vitro. A micellar solution containing 1 mmol/L oleic acid (Sigma),
0.5 mmol/L 1-monooleoylglycerol (Sigma), 6.6 mmol/L sodium tau-
rocholate, 0.6 mmol/L egg phosphatidylcholine (Sigma), and 132
mmol/L sodium chloride in 15 mmol/L sodium phosphate buffer (pH
6.8) was prepared by sonication and was kept at 37C for 24 h for
stabilization of the micelles. A solution of tea catechins or heat-
treated tea catechins was added to the micelles (final concentration,
2 g/L micelles) and incubated for 1 h at 37C. The micellar solution
was passed through a 0.2- m syringe filter (25 mm, GDD/X; What-
man). After lipid extraction from the filtrate, the concentration of
total fatty acids was measured by GLC (24).
Activity of pancreatic lipase in vitro. The activity of pancreatic
lipase was measured according to the method of Han et al. (18). An
emulsion (9 mL) containing 80 mg trioleoylglycerol, 10 mg phos-
phatidylcholine, and 5 mg sodium taurocholate in 0.1 mol/L
N-tris(hydroxymethyl)methyl-2-aminoethane sulfonic acid (TES)
buffer (pH 7.0) containing 0.1 mol/L sodium chloride was prepared by
sonication and kept at 37C. A total of 100 L of the emulsion was
incubated with 5 U of porcine pancreatic lipase (Sigma) solubilized in
0.1 mol/L TES buffer containing 0.1 mol/L sodium chloride and
various amounts of catechin solution (100 L) at 37C for 30 min.
Released fatty acids were extracted with chloroform: heptane: meth-
anol (49:49:2, v:v:v) and colorimetrically measured.
Statistical analysis. Data are expressed as means SEM. Sta-
tistical analysis of data were performed by two-way ANOVA or
two-way repeated-measure ANOVA followed by the Tukey-Kramer
test to identify differences among groups. Differences were considered
significant at P 0.05.
RESULTS
Postprandial hypertriacylglycerolemia in rats. Serum tri-
acylglycerol concentration increased after the administration
of a fat emulsion and was highest at 2 h after administration
(Fig. 2). The serum triacylglycerol concentration at 1, 2, and
TABLE 1
Composition of tea catechins and heat-treated tea catechins1
Catechins Tea catechins Heat-treated tea catechins
g/100 g dry weight
EGCG 34.2 19.7
GCG 5.8 20.3
ECG 16.4 10.8
CG 1.9 8.2
EGC 0.3 0.2
GC nd 0.1
EC 0.6 0.5
( )C 0.3 nd
Total 59.6 59.8
Gallate esters 58.3 59.0
Caffein nd nd
1 nd, not detected.
 TEA CATECHINS AND HYPERTRIACYLGLYCEROLEMIA
FIGURE 2 Effect of tea catechins and heat-treated tea catechins
on postprandial hypertriacylglycerolemia in rats intragastrically admin-
istered fat emulsions. Values are means SEM, n 7. Means not
sharing a letter at a time point differ, P 0.05. *Within a group, means
differ from that at time 0, P 0.05. Two-way repeated-measure
ANOVA: effect of group, P 0.075; effect of time, P 0.0001; inter-
action between group and time, P 0.0001.
3 h after administration was significantly lower in the tea
catechin and heat-treated tea catechin groups than in the
control group. There was no difference between the 2 cate-
chin-treated groups. 14
Lymphatic recovery of C-trioleoylglycerol in rats cannu-
lated in the thoracic14 duct. Lymphatic recoveries of the
radioactivity given as C-trioleoylglycerol at 1 h after admin-
istration were significantly lower in the tea catechin and
heat-treated tea catechin groups than in the control group
(Fig. 3). The tea catechins and heat-treated tea catechins had
similar effects on the lymphatic absorption of trioleoylglycerol.
Total 24-h recovery of the radioactivity was the same among
the 3 groups.
Micellar solubility of hydrolysis products of
triacylglycerol in vitro. When bile-salt micelles containing
oleic acid, 1-mono- oleoylglycerol, and phosphatidylcholine
were incubated with tea catechins or heat-treated tea
catechins in vitro, no precip- itates were observed, and the
concentrations of total fatty acids originating from oleic
acid, monooleoylglycerol, and phosphatidylcholine in bile-
salt micelles were not affected by the addition of tea
catechins or heat-treated tea catechins (data not shown).
Activity of pancreatic lipase in vitro. The activity of
pancreatic lipase was dose-dependently inhibited by the
addition of tea catechins and heat-treated tea catechins
(Fig. 4). There was no difference between the 2 catechin
preparations.
Catechins with a galloyl moiety dose-dependently inhibited
the activity of pancreatic lipase (Fig. 5). CG and GCG more
effectively inhibited the lipase activity than ECG and EGCG
at 1 and 2 mmol/L. Catechins without a galloyl moiety, C, EC,
GC, and EGC did not inhibit pancreatic lipase activity (data
not shown).
158
IKEDA ET AL.
157
DISCUSSION
Our results showed, for the first time, that tea catechins and
heat-treated tea catechins suppressed postprandial hypertriac-
ylglycerolemia to almost the same extent in rats (Fig. 2).
Because lymphatic recovery of trioleoylglycerol was delayed
by the ingestion of tea catechins and heat-treated tea
catechins (Fig. 3), one cause of the suppression of
postprandial hyper- triacylglycerolemia by the catechin
preparations may be the slower absorption of triacylglycerol
in the intestine. Both tea catechins and heat-treated tea
catechins dose-dependently inhibited the activity of pancreatic
lipase in vitro (Fig. 4). The results strongly suggest that the
delayed transport of trioleoyl- glycerol to the lymph is caused
by the inhibition of pancreatic lipase.
In the study of suppression of postprandial hypertriacyl-
glycerolemia (Fig. 2), catechins were administered at a con-
centration of 30 mg/g trioleoylglycerol. If we assume that one
human meal contains 1530 g of fats, 450 900 mg of cat- D
echins may be effective doses to inhibit postprandial hyper- o
triacylglycerolemia in humans. Green tea beverages in Japan w
contain 250 540 mg of heat-treated tea catechins per nl
serving. Therefore, ingestion of a sufficient amount of oa
catechins is possible. In a preliminary human study, we de
observed that postprandial hypertriacylglycerolemia in d
normal subjects was suppressed by the administration of a tea fro
beverage containing about 700 mg/d of heat-treated m jn.
catechins (unpublished re- sults). nu
In our previous study, the effect of tea catechins rich in trit
EGCG and ECG on lymphatic transport of fatty acids after io
intragastric administration of coconut oil or palm olein was n.
examined in rats cannulated in the thoracic duct (11). When or
coconut oil was given, tea catechins delayed lymphatic recov- g
ery of fatty acids and the result was consistent with the at
observation in this study. When palm olein was given, there Ku
was no delayed lymphatic recovery, but total recovery for 24 h nmi
of fatty acids was lower by the administration of tea catechins. ng
Therefore, the effect of tea catechins on fat absorption was In
sti
tut
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of
Bo
ta
ny,
C
A
S
on
Fe
br
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FIGURE 3 Effect of tea catechins and heat-treated tea catechins
on lymphatic recovery of [4- 14C] trioleoylglycerol in rats intragastrically
administered fat emulsions. Values are means SEM, n 7. Means
not sharing a letter at a time differ, P 0.05. Two-way repeated-
measure ANOVA: effect of group, P 0.14; effect of time, P 0.0001;
interaction between group and time, P 0.46.

FIGURE 4 Effect of tea catechins and heat-treated tea catechins
on the activity of pancreatic lipase in vitro. The enzyme activity at 0 g/L
was considered 100%. Values are means SEM, n 3. Within a
catechin preparation, means not sharing a letter differ, P 0.05.
Two-way ANOVA: effect of type of catechin, P 0.11; effect of cate-
chin concentration, P 0.0001; interaction between type of catechin
and catechin concentration, P 0.17.
equivocal in our previous work (11). In that study, lymphatic
recovery of fatty acids was analyzed by GLC. Because endog-
enous fatty acids were contained in lymph fluids, quantitative
estimation of absorbed fatty acids was impossible. In the
present study, because lymphatic recovery of fatty acids was
quantified using radiolabeled trioleoylglycerol, we think that
precise information on the effect of tea catechins on fat
absorption was obtained.
Han et al. (25) showed that tea saponin affected pancreatic
lipase activity in vitro. However, they did not report the data,
indicating that tea catechins did not inhibit the activity of
pancreatic lipase. Although we followed their method to mea-
sure the inhibitory effect of tea catechins on pancreatic lipase
activity, both tea catechins and heat-treated tea catechins
dose-dependently inhibited pancreatic lipase activity (Figs. 4
and 5). Causes of the discrepancy are not clearly explained at
present. It is also not evident if tea catechins and heat-treated
tea catechins inhibit pancreatic lipase activity in vivo. How-
ever, because our study simultaneously showed the
suppression of postprandial hypertriacylglycerolemia and
delayed recovery of triacylglycerol in lymph by these catechin
preparations, it is possible that tea catechins and heat-
treated tea catechins inhibit pancreatic lipase activity.
Both tea catechins and heat-treated tea catechins contain
unidentified components other than catechins (Table 1),
likely polymerized or degradated catechins. It is possible that
unknown components in catechin preparations also influence
the activity of pancreatic lipase. In the present study, major
components of heat-treated tea catechins GCG and CG more
effectively inhibited the lipase activity than those of tea cat-
echins EGCG and ECG (Fig. 5). Although these results sug-
gest that heat-treated tea catechins that are rich in GCG and
CG may more effectively inhibit pancreatic lipase than tea
catechins that are rich in EGCG and ECG, different inhibi-
tory effects on the lipase activity were not observed between
tea catechins and heat-treated tea catechins (Fig. 4). We also
showed that inhibitory effects by tea catechins and heat-
treated tea catechins on postprandial hypertriacylglycerolemia
and on lymphatic absorption of triacylglycerol were compara-
ble (Figs. 2 and 3). These results suggest that unidentified
components contained in the preparations of tea catechins
and heat-treated tea catechins may partly contribute to the
inhibition of pancreatic lipase.
The activation of lipoprotein lipase may be another impor-
tant determinant of postprandial triacylglycerol concentration
in serum. However, because studies have never been con-
ducted on this point, more studies on the effect of tea cat-
echins on the activity of lipoprotein lipase are required.
Our previous study showed that tea catechins and heat-
treated tea catechins precipitated cholesterol from bile-salt
micellar solutions and inhibited cholesterol absorption in rats
(12). In the present study, the addition of these tea catechins
to a bile-salt micellar solution containing the hydrolysis prod-
ucts of triacylglycerols, fatty acids, and monooleoylglycerol did
not affect the total fatty acid content in the micelles. The
results show that the catechin preparations do not exclude
oleic acid and monooleoylglycerol from a bile-salt micellar D
solution, suggesting that catechin preparations do not suppress ow
triacylglycerol absorption by inhibiting micellar solubility of nl
hydrolysis products of triacylglycerols. oa
Murase et al. (6) showed that long-term feeding of tea de
catechins, the major component of which was EGCG, reduced d
the deposition of visceral fat in mice fed a high-fat diet. They fro
ascribed the reduced deposition of visceral fat to enhanced m
-oxidation in the liver. We suggest that the suppression of jn.
postprandial hypertriacylglycerolemia may be another reason nu
for the antiobesity activity of tea catechins. Han et al. (25,26) trit
io
have pointed out that slower absorption of dietary fat n.
decreases the deposition of visceral fat. After a meal, the or
increase in blood glucose stimulates the secretion of insulin. g
Insulin acti- vates peripheral lipoprotein lipase and then at
hydrolysis of chy- lomicron triacylglycerols is stimulated. Ku
Formed FFAs are mainly incorporated into adipose tissue n
and deposited as tri- mi
ng
In
sti
tut
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of
Bo
ta
ny,
C
A
S
on
Fe
br
ua
FIGURE 5 Effect of purified catechins with a galloyl moiety on the
activity of pancreatic lipase in vitro. The enzyme activity at 0 mmol/L is
100%. Values are means SEM, n 6. Means not sharing a letter at
a concentration or within a catechin differ, P 0.05. Two-way ANOVA:
effect of type of catechin, P 0.0001; effect of catechin concentration,
P 0.0001; interaction between type of catechin and catechin con-
centration, P 0.0001.
 TEA CATECHINS AND HYPERTRIACYLGLYCEROLEMIA
acylglycerols. If the concentration of chylomicron triacylglyc-
erols is higher, it is possible that the deposition of fatty acids
hydrolyzed from triacylglycerols can be higher in adipose tis-
sues. Therefore the suppression of postprandial hypertriacyl-
glycerolemia by tea catechins may be a cause of the reduction
of fat deposition.
In conclusion, our study suggests that both green tea cat-
echins and heat-treated tea catechins suppress postprandial
hypertriacylglycerolemia, a risk factor for the development of
coronary heart disease. Our results, together with previous
observations, such as cholesterol-lowering activity (27,28) and
prevention of LDL oxidation in vitro (13) and in human
studies (14,15), strongly suggest that the ingestion of tea
catechins may prevent coronary heart disease. Our results also
suggest that heat-treated tea catechins contained in canned
and bottled tea beverages widely consumed in Asian countries
have the same effect on postprandial hypertriacylglycerolemia
as tea catechins.
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