Nutrient Metabolism

␤-Glucan Fractions from Barley and Oats Are Similarly Antiatherogenic

in Hypercholesterolemic Syrian Golden Hamsters1
Bryan Delaney, Robert J. Nicolosi,*2 Thomas A. Wilson,* Ting Carlson,† Scott Frazer,
Guo-Hua Zheng,† Richard Hess, Karen Ostergren, James Haworth and Nathan Knutson
Cargill Health and Food Technologies, Wayzata, MN 55391; *Center for Health and Disease Research,
Department of Health and Clinical Sciences, University of Massachusetts-Lowell, Lowell, MA 01854;
and †Cargill Sweeteners North America, Dayton, OH 45414-4321

ABSTRACT The cholesterol-lowering activities of oats and barley are commonly attributed to the ␤-glucan
fractions. Although ␤-glucan is present in both grains and appears to be chemically similar, the effect of source on
cholesterol-lowering activity has not been evaluated. In the present study, the antiatherogenic properties of
␤-glucan concentrates from oats and barley were evaluated in Syrian golden F1B hamsters consuming a semi-
purified hypercholesterolemic diet (HCD) containing cholesterol (0.15 g/100 g), hydrogenated coconut oil (20 g/100
g) and cellulose (15 g/100 g). After a 2-wk lead-in period, control hamsters were fed the HCD, whereas experi-
mental hamsters consumed HCD formulated to include ␤-glucan (2, 4, or 8 g/100 g) by addition of ␤-glucan

Downloaded from jn.nutrition.org by on January 27, 2009

concentrate prepared from oats or barley at the expense of cellulose. Compared with control hamsters, dose-
dependent decreases that were similar in magnitude in plasma total and LDL cholesterol concentrations were
observed in hamsters fed ␤-glucan from either source at wk 3, 6 and 9. Compared with controls, liver cholesterol
concentrations were also reduced (P ⬍ 0.05) in hamsters consuming 8 g/100 g oat or barley ␤-glucan. In agreement
with previously proposed mechanisms, total fecal neutral sterol concentrations were significantly increased (P
⬍ 0.05) in hamsters consuming 8 g/100 g barley or oat ␤-glucan. Aortic cholesterol ester concentrations were
significantly reduced (P ⬍ 0.05) in hamsters fed 8 g/100 g ␤-glucan from barley or oats. Although aortic total
cholesterol and cholesterol ester concentrations were significantly correlated with LDL cholesterol (r ⫽ 0.565, P
⬍ 0.004 and r ⫽ 0.706, P ⬍ 0.0001, respectively), this association could explain only half of the variability. This
study demonstrated that the cholesterol-lowering potency of ␤-glucan is approximately identical whether its origin
was oats or barley. J. Nutr. 133: 468 – 495, 2003.

KEY WORDS: ● ␤-glucan ● LDL cholesterol ● fecal neutral sterol ● aortic cholesterol ester
● early atherosclerosis

Consumption of oats or oat bran lowers serum cholesterol
concentrations in animal models of hypercholesterolemia (1–
5). The relevance of these studies to humans has been con-
firmed by numerous clinical studies demonstrating cholesterol-
lowering activity in individuals with elevated serum
cholesterol concentrations after consumption of oats (6,7) or
oat bran [(8 –11); reviewed in (12)]. This activity is largely
attributable to the soluble fiber fraction of oats, in particular to
the (133,134)-␤-D-glucan (␤-glucan) component. Because
the actual concentration of ␤-glucan in oats is relatively small
[⬍5%; 13–15], clinically relevant reductions in serum choles-
terol require the consumption of large amounts of whole-grain
foods (16). Alternatively, ␤-glucan concentrates have been
prepared from oats to enable oral consumption of ␤-glucan in
amounts likely to be associated with health benefits while
decreasing the need to consume large amounts of whole grains.
1
This study was sponsored by Cargill, Inc., Health and Food Technologies,
Wayzata, MN.
2
To whom correspondence should be addressed.
E-mail: Robert_Nicolosi@uml.edu.
Such preparations have demonstrated cholesterol-lowering ac-
tivity in hypercholesterolemic hamsters (1–3,17) and humans
(16,18).
Although fewer clinical studies exist, barley foods also
lower serum cholesterol concentrations in humans (19 –23).
As with oats, domestic barley cultivars contain ␤-glucan,
although at higher concentrations than in oats (15,24). How-
ever, although barley cultivars with the highest concentrations
of ␤-glucan contain approximately twice the amount found in
oats, consumption of large amounts of barley foods is also
likely to be necessary for clinically relevant reductions in
serum cholesterol concentrations. As with oats, concentrated
␤-glucan preparations from barley lower serum cholesterol
concentrations in animal models of hypercholesterolemia in-
cluding hamsters (2,25,26) and rats (27).
Modest differences may exist, but the ␤-glucan in oats and
barley appears to be similar structurally (28). It is therefore
likely that ␤-glucan concentrates from either source would
possess similar cholesterol-lowering activities. The present
study was designed to compare the effects of concentrated
␤-glucan from oats and barley on plasma lipids and lipoprotein

0022-3166/03 $3.00 © 2003 American Society for Nutritional Sciences.

Manuscript received 28 August 2002. Initial review completed 25 September 2002. Revision accepted 5 November 2002.

468
 CHOLESTEROL LOWERING BY BARLEY AND OAT ␤-GLUCAN
and hepatic cholesterol, fecal excretion of neutral sterols and
total bile acids, and antiatherogenic properties in hamsters
consuming a hypercholesterolemic diet (HCD)3. Hamsters
were selected for this study because decreased plasma choles-
terol concentrations have been reported in hamsters fed HCD
after consumption of ␤-glucan– enriched fractions from oats
and barley (1– 4,17,25). Furthermore, hamsters represent a
useful model for humans because they metabolize cholesterol
similarly (29,30) and their lipoprotein profile resembles that of
humans fed HCD (2). The relevance of the hamster model to
humans has been established by the consistent ability of ham-
sters to respond to dietary modulation of cholesterol absorp-
tion similarly to humans (2,30).
MATERIALS AND METHODS
Preparation of ␤-glucan concentrate from barley and oats.
␤-Glucan concentrate (Barley Betafiber, Cargill, Inc., Wayzata, MN)
was prepared from hulless barley (Azhul variety) by an extraction process
similar to that of Aman and Hesselman (31) with slight modifications.
Ground barley flour was mixed with boiling water and bacterial ␣-amy-
lase to extensively hydrolyze starch, solubilize ␤-glucan and inactivate
contaminating ␤-glucanase activities. Solubilized ␤-glucan in the clari-
fied liquid fraction was precipitated, washed with ethanol and dried. A
similar procedure was used for the extraction of ␤-glucan from oat bran
(SCM 350 oat bran from ConAgra Foods, S. Sioux City, NE) except
protease (papain) was used to hydrolyze protein in addition to the
starch hydrolysis step and hexane was used to extract excess oil from
dried oat ␤-glucan concentrate. Proximate analysis of the concen-
trated ␤-glucan preparations is presented in Table 1.
Diets. All diets were formulated and dietary ingredients supplied
by Research Diets (New Brunswick, NJ) except for barley and oat
␤-glucan concentrates, which were prepared by Cargill, Health and
Food Technologies (Wayzata, MN). Diet ingredients for all groups
were identical except for the nutrient under evaluation as indicated
in Table 2. The control HCD contained 15 g/100 g cellulose, whereas
the experimental diets were formulated to contain 2, 4 or 8 g/100 g
␤-glucan from oats or barley by the addition of the ␤-glucan concen-
trate from the respective source at the expense of cellulose. Analyt-
ical characterization of ␤-glucan was conducted on hamster feed to
validate the composition and homogeneity of dietary blending and
stability under conditions of storage and use. Hamster feed was milled
and ␤-glucan was quantified using a Megazyme (Megazyme Interna-
tional Ireland, Bray, Wicklow, Ireland) kit version of the McCleary
method [AOAC method 995.16; see (32,33)].
Animals. Male Syrian golden hamsters (n ⫽ 70; F1B strain,
BioBreeders, Fitchburg, MA) ⬃8 –10 wk old were acclimated to
individual stainless steel cages and fed HCD for 2 wk before initiation
of the experimental diets. Hamsters were then bled to determine
plasma concentrations of total (TC), HDL (HDL-C) and LDL cho-
lesterol (LDL-C); they were divided into 7 groups (n ⫽ 10/group)
with similar starting plasma LDL-C concentrations and fed the spec-
ified diets. The experimental protocols were approved by the Insti-
tutional Animal Care and Use Committee. Hamsters were main-
tained in accordance with guidelines of the Animal Care Committee
at the University of Massachusetts-Lowell Research Foundation and
the NIH. Hamsters were housed in environmentally controlled con-
ditions with an alternating 12-h light:dark cycle and consumed food
and water ad libitum except when food was withheld for the exper-
imental protocols described below.
Plasma lipoprotein cholesterol and triglyceride measurements.
Blood was collected via the retro-orbital sinus into heparinized tubes
from hamsters deprived of food for 12 h. Plasma was harvested after
centrifugation at 1500 ⫻ g at room temperature for 10 min, and
plasma TC (34) and triglyceride (TG) (35) concentrations were
measured enzymatically. Plasma LDL-C (combination of VLDL, in-
3
Abbreviations used: GC, gas chromatography; HCD, hypercholesterolemic
diet; HDL-C, HDL cholesterol; LDL-C, LDL cholesterol; TC, total cholesterol; TG,
triglyceride.
469
TABLE 1
Proximate composition of ␤-glucan concentrates
from barley and oats
%
Barley Oats
␤-Glucan1 77.98 64.88
Hydrolyzed starch2 6.53 8.66
Protein3 7.85 11.10
Moisture4 5.20 8.80
Oil5 0.96 1.64
Ash6 1.82 2.78
1 ␤-Glucan concentration was determined by AOAC method 995.16
using Megazyme mixed-linkage ␤-glucan assay kit (32, 33).
2 Hydrolyzed starch was determined enzymatically using amyloglu-
cosidase.
3 Protein concentration was determined by AOAC method 991.20
(32).
4 Moisture was determined by AOAC method 926.08 (32).
5 Oil concentration was determined by AOAC method 933.05 (32).
6 Ash concentration was determined using Corn Refiners Associa-
Downloaded from jn.nutrition.org by on January 27, 2009
tion Standard Analytical Method A-4 (62).
termediate, and LDL-C) was precipitated with phosphotungstate
reagent (36) and HDL-C was measured in the supernatant. The
concentration of LDL-C was calculated as the difference between
plasma TC and HDL-C. The accuracy and precision of the procedures
used for the measurements of plasma TC and HDL-C were main-
tained by participation in the Lipid Standardization Program of the
Centers for Disease Control and the National Heart, Blood and Lung
Institute (Atlanta, GA). All reagents used were supplied by Sigma-
Aldrich (St. Louis, MO). All assays were performed using a Cobas
Mira Plus (Roche Diagnostic Systems, Basel, Switzerland) clinical
chemistry autoanalyzer using 10 ␮L of plasma for each sample (Roche
Pharmaceuticals).
Aortic cholesterol measurements. At the end of the exposure
period (wk 9), hamsters were anesthetized with an intraperitoneal
injection of sodium pentobarbital and aortic tissue was obtained for
determination of cholesterol concentration. The heart and thoracic
aorta were removed and stored in PBS at 4°C for subsequent analysis.
To measure cholesterol concentrations in the aortic arch, a piece of
aortic tissue extending from as close to the heart as possible to the
branch of the left subclavian artery was used (⬃20 – 40 mg). The
tissue was cleaned, weighed and placed in a vial containing 4 mL of
methanol and 10 mL of chloroform and treated as described by Rudel
et al. (37). The sample was mixed vigorously and left at room
temperature for 48 h before extraction. The solution was then placed
in a 37°C water bath, under N2. When one half of the solution was
evaporated, 1 mL of chloroform with 1% Triton-100 was added,
mixed and evaporated to dryness at 37°C under N2. Distilled water
(250 ␮L) was added to the samples, vortexed and placed in a shaking
water bath at 37°C for 20 min to solubilize the lipid. After incuba-
tion, aortic total and free cholesterol concentrations were determined
enzymatically in triplicate with 25 ␮L of sample (Wako Chemicals,
Richmond, VA) using an ELISA assay. The aortic cholesteryl ester
concentration was calculated as the difference between the total and
the free cholesterol concentrations. A pilot study was conducted to
evaluate the extent to which this procedure removed tissue choles-
terol. Aortic cholesterol concentrations were determined after tissue
was placed in solvent (4 mL of methanol and 10 mL of chloroform)
overnight with frequent vigorous mixing and compared with the
concentrations obtained after tissue minced or homogenization. No
significant differences in aortic free or cholesterol ester content were
observed between the different cholesterol extraction procedures
with efficiencies of ⬃96%.
Hepatic cholesterol measurements. Hepatic cholesterol concen-
trations were measured by a previously described method (38) in the
following manner: a 100-mg portion of liver was homogenized with
470 DELANEY ET AL.

TABLE 2
Diet composition

Barley ␤-glucan (g/100 g) Oat ␤-glucan (g/100 g)

Control 2 4 8 2 4 8

g/kg

Casein, 80 mesh 250 250 250 250 250 250 250

DL-Methionine 5 5 5 5 5 5 5
MaltoDextrin 10 125 125 125 125 125 125 125
Cornstarch 156 156 156 156 156 156 156
Sucrose 95 95 95 95 95 95 95
Cellulose BW200 150 121.5 93 36 116.75 83.5 17
Barley ␤-glucan concentrate1 0 28.5 57 114 0 0 0
Oat ␤-glucan concentrate 0 0 0 0 33.25 66.5 133
Safflower oil 20 20 20 20 20 20 20
Coconut oil, hydrogenated 200 200 200 200 200 200 200
Mineral mix, S100012 35 35 35 35 35 35 35
Vitamin mix, V100013 10 10 10 10 10 10 10
Choline bitartrate 2 2 2 2 2 2 2
Cholesterol 1.5 1.5 1.5 1.5 1.5 1.5 1.5
Yellow dye, FD&C#5 0 0.1 0 0 0.05 0.05 0
Red dye, FD&C#40 0 0 0.1 0 0.05 0 0.05

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Blue dye, FD&C#1 0 0 0 0.1 0 0.05 0.05

1 Barley Betafiber, Cargill, Inc., Wayzata, MN

2 Mineral mix, S10001 composition (g/kg): calcium carbonate, 160.0; calcium phosphate, 235.0; magnesium oxide, 5.0 g; potassium citrate, 420.0
g; potassium sulfate, 20.0; sodium chloride, 66.7; chromium potassium sulfate 0.1; cupric carbonate, 0.14; potassium iodate, 0.02; ferric citrate, 4.0;
manganous carbonate, 1.4; sodium selenite, 0.003; zinc carbonate, 0.75; sucrose, 46.887.
3 Vitamin mix, V10001 composition (amount in 10 grams): vitamin A palmitate, 20,000 IU; vitamin D-3, 1,000 IU; vitamin E acetate, 50 IU; menadione
sodium bisulfate, 0.5 mg; biotin, 0.3 mg; cyanocobalamin, 10 ␮g; folic acid, 6 mg; nicotinic acid, 30 mg; calcium pantothenate, 30 mg; pyridoxine-HCl,
6 mg; riboflavin, 6 mg; thiamin-HCl, 6 mg; ascorbic acid, 500 mg.

50 mg of sodium sulfate. Methanol (5 mL) was then added and the
tissue homogenized a second time followed by addition of 10 mL of
chloroform. After mixing, 3 mL of a solution containing 1.25% KCl
and 0.05% H2SO4 was added and centrifuged at 400 ⫻ g at room
temperature for 10 min. The bottom layer was transferred and the
supernatant reextracted with 3 mL of chloroform/methanol (2:1) and
centrifuged at 400 ⫻ g at room temperature for 10 min. The bottom
layer was transferred and pooled with the previous step. The solution
was placed in a 37°C water-bath and placed under N2. When ap-
proximately half of the solution was evaporated, 1 mL of chloroform
with 1% Triton-100 was added, mixed and evaporated to dryness at
37°C under N2. Distilled water (500 ␮L) was added to the samples,
vortexed and placed in a shaking water bath at 37°C for 20 min to
solubilize the lipid. After incubation, hepatic total and free choles-
terol concentrations were determined enzymatically (Wako Chemi-
cals) using an ELISA assay. Hepatic cholesteryl ester concentration
was calculated as the difference between the total and the free
cholesterol concentrations.
Fecal neutral sterol measurements. Fecal samples were col-
lected over the final 3 d of the exposure period, freeze-dried (lyoph-
ilized) and ground before analysis (39). Dry feces (200 mg) were
extracted with 4 mL of methanol/chloroform (50:50) for 1 h at 100°C
in a 5-mL Reacti-vial (Pierce, Rockford, IL) fitted with a mini-nert
cap. Samples were then allowed to come to room temperature and
were centrifuged at 500 ⫻ g at room temperature for 10 min. The
supernatant was removed from the fecal pellet and transferred to an
8-mL borosilicate vial. Supernatants were evaporated to dryness at
50°C under N2. Then, 4 mL of 0.1 mol/L NaOH/ethanol (10:90, v/v)
was added to each sample, overlaid with N2, capped and heated at
100°C for 30 min. The samples were allowed to cool to room
temperature; the solvent was removed and transferred to 16 ⫻ 150
mm borosilicate test tubes. Water (5 mL) and 3 mL of hexane were
added to the solvent followed by vortexing and centrifugation at 500
⫻ g for 2 min. The top hexane layer was removed and placed in vials.
The hexane extraction was repeated two more times and pooled. The
hexane extracts were stored at ⫺80°C until analysis of neutral sterols.
The aqueous layer was adjusted to 10 mL with the addition of 0.1
mol/L NaOH, and 1 mL was removed for quantitative enzymatic
determination of total bile acids. To the hexane portion, 1 mL of
5-␣-cholestane (240 mg/L) (Sigma-Aldrich) was added and the so-
lution was adjusted to 10 mL with hexane in a volumetric flask. After
2 mL were removed and evaporated to dryness at 100°C under N2,
100 ␮L of Tri-Sil reagent was added and the samples heated at 85°C
for 20 min, followed by evaporation and reconstitution in 100 ␮L of
methylene chloride. Then 1 ␮L was injected and analyzed by capil-
lary gas chromatography (GC).
GC analyses. Neutral sterols were analyzed using a Shimadzu
(Kyoto, Japan) GC-14A gas chromatograph with a flame ionization
detector and a 50 m ⫻ 0.2 mm HP-1 capillary column (Hewlett
Packard, Andover, MA). The injector and detector temperatures
were set at 300°C. The initial column temperature was 220°C and
was increased to 300°C at a rate of 2°C/min. The final temperature
was held for 10 min. Column flow rate was 1.5 mL/min. Peak areas
were quantitated using a Shimadzu CR501 integrator. Quantification
and identification of neutral sterols were based on the appropriate
purified external standards supplied by Sigma-Aldrich. Extraction
efficiency for neutral sterol sterols following this protocol was ⬃94%.
Total fecal bile acid measurements. Using Sigma Procedure
#450, blank and test reagents were prepared as described in assay
instructions. Standard solutions were prepared using known quanti-
ties of lithocholic acid (Sigma-Aldrich). Standards and samples (16
␮L of the 1 mL aqueous portion plus 24 ␮L of calf serum) were
assayed in two sets of triplicates (one set for 100 ␮L of test reagent,
the other set for 100 ␮L of blank reagent) on 96-well microtiter plate.
After 10 min incubation at 37°C, absorbances were read at 530 nm
using a Molecular Devices (Sunnyvale, CA) microplate reader. The
extraction efficiency for bile acids from hamster feces following this
protocol is ⬃92%.
Statistical analysis. One-way ANOVA was used to examine the
effect of treatment on the different variables using SigmaStat software
(Jandel Scientific, San Rafael, CA). Plasma lipid and lipoprotein
cholesterol concentrations in ␤-glucan–treated hamsters were com-
pared with the HCD control group using Dunnett’s t test. Intergroup
comparisons (e.g., 2 g/100 g oats vs. 2 g/100 g barley) were evaluated
 CHOLESTEROL LOWERING BY BARLEY AND OAT ␤-GLUCAN 471

TABLE 3
Plasma total cholesterol concentrations of hamsters consuming concentrated ␤-glucan from oats or barley1

␤-Glucan concentrate wk 0 wk 3 wk 6 wk 9

mmol/L

0 (Control 5.86 ⫾ 0.47a 7.61 ⫾ 1.13a 7.43 ⫾ 0.56a 7.60 ⫾ 0.86a

Oat, 2 g/100 g 6.02 ⫾ 0.69a 6.46 ⫾ 1.78a 7.75 ⫾ 0.63a 7.97 ⫾ 1.08a
Barley, 2 g/100 g 5.84 ⫾ 0.61a 7.51 ⫾ 1.08a,2 7.43 ⫾ 0.49a 6.97 ⫾ 1.02a
Oat, 4 g/100 g 5.72 ⫾ 0.54a,2 5.65 ⫾ 1.65b,c 6.24 ⫾ 0.70b,c 6.33 ⫾ 1.07b,c
Barley, 4 g/100 g 5.80 ⫾ 0.58a 6.06 ⫾ 1.32b,3 6.49 ⫾ 0.90b 6.53 ⫾ 0.98b
Oat, 8 g/100 g 6.09 ⫾ 0.84a 4.95 ⫾ 0.54c 5.52 ⫾ 0.61c 5.15 ⫾ 0.83c
Barley, 8 g/100 g 6.12 ⫾ 0.84a 5.24 ⫾ 0.83c 5.85 ⫾ 1.00c 5.58 ⫾ 1.09c

1 Values are means ⫾ SD, n ⫽ 10 except where noted. Means in each column with different superscript letters differ (P ⬍ 0.05).
2 n ⫽ 9.
3 n ⫽ 8.

using Student’s t test. Correlations between LDL-C and aortic TC consuming 2 g/100 g ␤-glucan from either source. The de-
and aortic cholesterol ester were determined using pooled samples of crease in plasma TC concentrations was not distinguishable on
both 8 g/100 g ␤-glucan from barley and oats by Pearson’s Product- the basis of the source of ␤-glucan. There were no effects on
moment method as previously reported (40) and plotted using Sig- plasma TG concentrations, food intake and body weight in

Downloaded from jn.nutrition.org by on January 27, 2009

maStat software (Jandel Scientific, San Rafael, CA).
RESULTS
Isolation of ␤-glucan and dietary incorporation. Concen-
trations of ␤-glucan from the barley and oat preparations were
78 and 65%, respectively (Table 1). The ␤-glucan– enriched
fraction of oats was prepared using a similar process but yielded
65% ␤-glucan in the oat extract (Table 1). Because the
concentration of ␤-glucan from the barley concentrate was
higher than that of the oat preparation, the addition of smaller
amounts of the barley preparation was necessary to obtain the
same dietary concentration of ␤-glucan when incorporated
into the hamster feed. These fractions were then blended into
a semisynthetic HCD at concentrations corresponding to 2, 4,
or 8 g/100 g (Table 2). The ␤-glucan was incorporated homo-
geneously and was stable under the conditions used to blend,
ship and store the experimental diets (data not shown).
Plasma TC and TG concentrations. Plasma TC concen-
trations in hamsters consuming the HCD were approximately
the same in all groups after a 2-wk lead-in period (Table 3).
After that period, hamsters were fed the indicated diets. Com-
pared with hamsters fed the HCD, a dose-dependent decrease
in plasma TC concentrations was observed at all time points
after consumption of the experimental diets containing 4 and
8 g/100 g ␤-glucan from oats or barley, but not in hamsters
any of the treatment periods (data not shown).
Plasma LDL-C concentrations. Similar to plasma TC
concentrations, a dose-dependent decrease was observed in
LDL-C concentrations in hamsters fed diets supplemented
with 4 and 8 g/100 g ␤-glucan from either oats or barley (Table
4). However, at 2 g/100 g ␤-glucan from either source, no
changes were observed compared with HCD control. Impor-
tantly, the plasma LDL-C concentrations in hamsters treated
with the 4 and 8 g/100 g concentrations of ␤-glucan from
either source were not different from each other at any time
point.
Plasma HDL-C concentrations. Plasma concentrations of
HDL-C were similar in all groups after consumption of the
HCD for 2 wk (Table 5). Compared with the HCD control
group, consumption of 8 g/100 g ␤-glucan–supplemented diets
from both sources caused decreases in plasma HDL-C concen-
trations at 6 and 9 wk. Surprisingly, a significant increase in
plasma HDL-C concentrations compared with HCD control
(P ⬍ 0.05) was observed in the 2 g/100 g oat group that did
not occur in the barley ␤-glucan group at 9 wk.
Liver cholesterol concentrations. After 9 wk, liver total,
free and esterified cholesterol concentrations were significantly
decreased in hamsters consuming 8 g/100 g ␤-glucan from oats
or barley compared with the control group. Although the
difference in liver TC was significant (P ⬍ 0.05) compared

TABLE 4
Plasma LDL cholesterol concentrations of hamsters consuming concentrated ␤-glucan from oats or barley1

␤-Glucan concentrate wk 0 wk 3 wk 6 wk 9

mmol/L

0 (Control) 3.77 ⫾ 0.47a,2 4.91 ⫾ 1.24a 4.63 ⫾ 0.48a 5.07 ⫾ 0.76a

Oat, 2 g/100 g 3.92 ⫾ 0.88a 4.16 ⫾ 1.96a 4.80 ⫾ 0.42a 5.13 ⫾ 0.58a
Barley, 2 g/100 g 3.64 ⫾ 0.67a 4.82 ⫾ 1.21a 4.99 ⫾ 0.51a 4.45 ⫾ 1.01a
Oat, 4 g/100 g 3.70 ⫾ 0.59a 3.17 ⫾ 1.70b,c 3.84 ⫾ 0.53b 3.75 ⫾ 0.82c
Barley, 4 g/100 g 3.68 ⫾ 0.63a,2 3.93 ⫾ 1.59a 3.96 ⫾ 0.82b 4.08 ⫾ 0.89b
Oat, 8 g/100 g 3.96 ⫾ 0.65a 2.74 ⫾ 0.44c 3.28 ⫾ 0.52b 3.01 ⫾ 0.56c
Barley, 8 g/100 g 3.89 ⫾ 0.88a 2.82 ⫾ 0.82c 3.59 ⫾ 0.84b 3.53 ⫾ 0.93c

1 Values are means ⫾ SD, n ⫽ 10 except where noted. Means in each column with different superscript letters differ (P ⬍ 0.05).
2 n ⫽ 9.
472 DELANEY ET AL.

TABLE 5
Plasma HDL cholesterol concentrations of hamsters consuming concentrated ␤-glucan from oats or barley1

␤-Glucan concentrate wk 0 wk 3 wk 6 wk 9

mmol/L

0 (Control) 2.29 ⫾ 0.59a 2.70 ⫾ 0.46a 2.51 ⫾ 0.33a 2.53 ⫾ 0.31a

Oat, 2 g/100 g 2.09 ⫾ 0.26a 2.31 ⫾ 0.54a 2.77 ⫾ 0.31a 3.09 ⫾ 0.54b,2
Barley, 2 g/100 g 2.20 ⫾ 0.14a 2.64 ⫾ 0.36a 2.44 ⫾ 0.16a 2.53 ⫾ 0.26a
Oat, 4 g/100 g 2.03 ⫾ 0.33a 2.42 ⫾ 0.16a,2 2.41 ⫾ 0.01a 2.59 ⫾ 0.34a
Barley, 4 g/100 g 2.22 ⫾ 0.43a 2.51 ⫾ 0.35a 2.53 ⫾ 0.24a 2.45 ⫾ 0.46a
Oat, 8 g/100 g 2.23 ⫾ 0.23a 2.47 ⫾ 0.14a 2.24 ⫾ 0.20b 2.14 ⫾ 0.36b
Barley, 8 g/100 g 2.06 ⫾ 0.26a,2 2.42 ⫾ 0.35a 2.26 ⫾ 0.26b 2.05 ⫾ 0.28b

1 Values are means ⫾ SD, n ⫽ 10 except where noted. Means in each column with different superscript letters differ (P ⬍ 0.05).
2 n ⫽ 9.

with hamsters fed the HCD, there were no significant differ-
ences between oat or barley glucan groups (Fig. 1). Hepatic
cholesterol concentrations were not evaluated in hamsters
consuming 2 or 4 g/100 g ␤-glucan from oats or barley.
Fecal neutral sterol and total bile acid concentrations.
centration was observed in both the oat (⫺71%) and barley
(⫺58%) groups (P ⬍ 0.05; Fig. 2). The difference in aortic
TC concentration between the oat and barley groups was not
significant (P ⫽ 0.206). Although the concentration of aortic
free cholesterol was not different from the cholesterol control

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Consumption of 8 g/100 g ␤-glucan from oats or barley signif-
icantly increased (P ⬍ 0.05) total fecal neutral sterol concen-
trations by 76 and 78%, respectively (Table 6) relative to the
HCD. Virtually all individual sterols analyzed were greater for
hamsters fed the barley and oat ␤-glucan compared with the
HCD group, with the greatest quantitative increases in excre-
tion observed for cholesterol (169 –218%) and coprostanol
(43–52%). Although some differences were noted between
fecal excretion of individual and total neutral sterols between
the barley and oat groups, none were significant (P ⬎ 0.05).
No significant differences were observed between any dietary
treatments for fecal excretion of total bile acids (Table 6).
Fecal neutral sterol and total fecal bile acid concentrations
were not evaluated in hamsters consuming 2 or 4 g/100 g
␤-glucan from oats or barley.
Aortic cholesterol concentrations. Compared with the
HCD control group, a significant decrease in aortic TC con-
group in either of the ␤-glucan groups, the aortic cholesterol
ester concentrations were profoundly decreased in hamsters
fed 8 g/100 g ␤-glucan from oats (⫺84%) or barley (⫺80%).
When data from both 8 g/100 g oat and barley ␤-glucan groups
were pooled, the aortic TC and cholesterol ester concentra-
tions were significantly correlated with LDL-C (r ⫽ 0.565, P
⬍ 0.004, r ⫽ 0.706, P ⬍ 0.0001, respectively). However, only
50% of the variability could be attributed to this association.
Aortic cholesterol concentrations were not evaluated in ham-
sters consuming 2 or 4 g/100 g ␤-glucan from oats or barley.
DISCUSSION
The cholesterol-lowering activity of oats and barley is be-
lieved to be attributable to the ␤-glucan in the soluble fiber
fraction of these cereal grains. ␤-Glucan is composed of high-
molecular-weight water-soluble cell-wall polysaccharides con-
sisting of (133,134)-␤-D-linked glucopyranosyl-monomers
(41). The ␤-glucan in both grains is similar in structure;

TABLE 6
Fecal neutral sterol and total bile acid concentrations of
hamsters consuming concentrated ␤-glucan
from oats or barley1

8 g/100 g ␤-Glucan

Control Oat Barley

mg/g of feces

Cholesterol 1.34 ⫾ 0.9a 4.24 ⫾ 2.9b 3.59 ⫾ 2.8b

Coprostanol 5.15 ⫾ 2.0a 7.37 ⫾ 2.2b 7.83 ⫾ 1.9b
Coprostanone 0.06 ⫾ 0.1a 0.05 ⫾ 0.1a 0.10 ⫾ 0.2a
Campesterol 0.81 ⫾ 0.3a 1.82 ⫾ 0.5b 1.71 ⫾ 0.5b
Stigmasterol 0.11 ⫾ 0.1a 0.24 ⫾ 0.1b 0.16 ⫾ 0.1a
Sitosterol 0.09 ⫾ 0.0a 0.30 ⫾ 0.2b 0.24 ⫾ 0.2a
Sitostanol 0.09 ⫾ 0.1a 0.25 ⫾ 0.2a 0.25 ⫾ 0.2a
Total neutral sterols 7.63 ⫾ 3.2a 14.27 ⫾ 4.4b 13.86 ⫾ 5.0b
FIGURE 1 Concentrations of cholesterol (total, free, and esteri- Total fecal cholesterol 6.54 ⫾ 2.8a 11.51 ⫾ 4.2b 11.66 ⫾ 3.6b
fied) in the livers of male Syrian golden F1B hamsters after 9 wk Total fecal bile acids 2.69 ⫾ 1.5a 3.38 ⫾ 1.0a 2.65 ⫾ 1.1a
consumption of diets formulated with cellulose control, or 8 g/100 g
␤-glucan from concentrated oat or barley ␤-glucan. Data are expressed 1 Values represent means ⫾ SD, n ⫽ 10. Means in each row with
as means ⫾ SD, n ⫽ 10. Bars with different letters differ (P ⬍ 0.05). different superscript letters differ (P ⬍ 0.05).
 CHOLESTEROL LOWERING BY BARLEY AND OAT ␤-GLUCAN
FIGURE 2 Concentrations of cholesterol (total, free, and esteri-
fied) in the aortas of male Syrian golden F1B hamsters after 9 wk
consumption of diets formulated with cellulose control, or 8 g/100 g
␤-glucan obtained from concentrated oat or barley ␤-glucan. Data are
expressed as mean ⫾ SD, n ⫽ 8/group because 2 outliers were de-
tected in each group. Bars with different letters differ (P ⬍ 0.05).
however, differences with respect to the ratio of (133) to
(134)-linkages (42,43), molecular weight (28) and possibly
solubility (15) have been reported. Concentrated ␤-glucan
from both sources lowers plasma cholesterol concentrations in
clinical studies (16,18) and in animal models of hypercholes-
terolemia (1–3,17,25–27), but a comparison of the effects of
␤-glucan from different sources has not been reported.
In the current study, ␤-glucan concentrates were prepared
from oats and barley and their antiatherogenic properties
evaluated in hypercholesterolemic hamsters. Consumption of
semipurified diets containing concentrated oat or barley ␤-glu-
can indicated a dose-dependent decrease in plasma TC con-
centrations regardless of the source of the ␤-glucan concen-
trate. The magnitude of the decrease was consistent with
previous studies evaluating the effect of ␤-glucan concentrates
from barley (2,25) and similar to (3) or slightly greater in
magnitude than the effect of oat bran and oat ␤-glucan con-
centrates in hamsters (1,2,17,44). This effect was primarily
attributable to decreased plasma LDL-C concentrations. De-
creased HDL-C concentrations were also observed in hamsters
consuming high concentrations of ␤-glucan from oats and
barley as has been reported in other studies evaluating the
effect of barley or oat preparations in hamsters (2,317,25). The
clinical relevance of the latter observation is questionable
because it does not appear to occur in humans (6 – 8,10,
11,16,19,21). However, the results from this study demon-
strate that the plasma cholesterol–lowering activity of ␤-glu-
can is approximately the same whether it is from barley or oats.
Similar observations have been reported in rats consuming
HCD supplemented with oat and barley gums (45).
The ability of soluble fiber, and specific components
therein, to lower serum cholesterol concentrations is believed
to occur through a combination of mechanisms. Consumption
of ␤-glucan inhibits absorption of cholesterol from the gut as
demonstrated by a significant increase in the excretion of fecal
cholesterol and neutral sterols (7,44,46). In the current study,
a large increase in the fecal excretion of neutral sterols and
cholesterol was observed after consumption of concentrated
473
oat or barley ␤-glucan. Lia and co-workers (47) reported that
oats caused a greater increase in sterol excretion than barley;
however, there was no difference between oats and barley in
the current report. This observation supports the concept that
the cholesterol-lowering properties of ␤-glucan are attribut-
able at least in part to inhibition of cholesterol absorption
from the gut. Fibers containing ␤-glucan have also been re-
ported to increase excretion of bile acids, suggesting a caus-
ative role in the lowering of plasma cholesterol concentrations
(7,11,48). However, this association is not consistent because
numerous studies have reported a cholesterol-lowering effect
in the plasma without alteration of fecal bile acid concentra-
tions (49 –52). Neither source of ␤-glucan concentrate altered
fecal bile acid concentrations in the current study. Alterna-
tively, consumption of ␤-glucan– containing fibers increases
fecal output (7,48). Therefore, similar concentrations of bile
acid relative to fecal weight may represent an increase in bulk
excretion of bile acids; however, fecal output was not evalu-
ated in the current study.
Although many studies have shown that consumption of
␤-glucan decreases plasma lipid and lipoprotein cholesterol
concentrations, the current study also demonstrated a striking
decrease in the concentration of aortic cholesterol. This effect
Downloaded from jn.nutrition.org by on January 27, 2009
was attributable almost entirely to the cholesterol ester con-
tents of the aorta, and again there was no difference in mag-
nitude between oats and barley. This observation is particu-
larly important because this form of tissue cholesterol is viewed
as the hallmark in fatty streak formation. This is the first time
that dietary ␤-glucan concentrates from any source have dem-
onstrated such an effect on the aorta. Equally surprising was
the finding that only 50% of the association could be ex-
plained by the effects of ␤-glucan on plasma LDL-C concen-
trations.
This suggests that additional antiatherogenic mechanism(s)
of action of ␤-glucan may occur outside of the gut. For exam-
ple, some studies have reported a decrease in hepatic 3-hy-
droxy-3-methylglutaryl-CoA reductase activity after consump-
tion of diets containing barley or oats (53). Alternatively,
changes may also occur at the level of the blood vessel wall
within the aorta. Oxidative stress (54) and inflammation (55)
have been implicated in the development of early and ad-
vanced atherosclerosis. Therefore, dietary antioxidants could
contribute to the effects observed in aortic cholesterol ester
concentrations in the current study. For example, recent stud-
ies in guinea pigs have suggested that consumption of certain
fibers may have antioxidant-like properties that are measurable
in circulating LDL-C (56). Numerous antioxidants have been
identified in both barley and oats (57–59) that may also have
been concentrated in the process of preparing the ␤-glucan
concentrates used in the current study. Further, substances not
associated with the soluble fiber fractions have been identified
in these grains and may contribute to the effect by inhibiting
endogenous cholesterol production (60,61). However, it is not
known whether the procedures used to concentrate the ␤-glu-
can fractions concentrate or dilute these substances and their
concentrations were not evaluated in either ␤-glucan prepa-
ration used in this study.
The results of the current study demonstrated that ␤-glucan
concentrates from barley and oats modify plasma cholesterol
concentrations and other indicators associated with athero-
genic progression in hamsters with similar potency and
through similar mechanisms of action. The effects were de-
pendent on dietary concentrations of ␤-glucan but no differ-
ences were observed between the barley and oat preparations.
Therefore, the structural differences between the ␤-glucan in
barley and oats (15,28,42,43) may not be important with
474 DELANEY ET AL.
regard to the physiologic effects that occur after their con-
sumption. These results support the cholesterol-lowering and
antiatherogenic activity of ␤-glucan concentrates from barley
and oats.
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