Molecular Aspects of Cereal Glucan Functionality Physical Properties Technological Applications and Physiological Effects 2007 Journal of Cereal Scien

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Journal of Cereal Science 46 (2007) 101118

www.elsevier.com/locate/jcs
Molecular aspects of cereal b-glucan functionality: Physical properties,

technological applications and physiological effects
A. Lazaridou, C.G. Biliaderis
Laboratory of Food Chemistry & Biochemistry, Department of Food Science and Technology, School of Agriculture, Aristotle University,
P.O. Box 256, Thessaloniki 541 24, Greece
Received 2 February 2007; received in revised form 7 May 2007; accepted 8 May 2007
Abstract
Cereals b-glucans are linear homopolysaccharides of consecutively linked (1-4)-b-D-glucosyl residues (i.e. oligomeric cellulose
segments) that are separated by single (1-3)-linkages. b-Glucans display all the functional properties of viscous and gel forming food
hydrocolloids combined with all the physiological properties of dietary bres. This review focuses on the relationships between the
molecularstructural characteristics of b-glucans and their physicochemical properties in aqueous dispersions and in food systems as well
as their physiological functions in the gastro-intestinal tract. The physical properties of b-glucans, such as solubility and rheological
behaviour in the solution and gel states, are controlled by their molecular features, such as their distribution of cellulosic oligomers, their
linkage pattern and their molecular weight as well as by temperature and concentration. The technological and nutritional functionality
of b-glucans is often related to their rheological behaviour. Incorporation of b-glucans into various products (bread, mufns, pasta,
noodles, salad dressings, beverages, soups, reduced-fat dairy and meat products) showed that attributes, such as breadmaking
performance, water binding and emulsion stabilising capacity, thickening ability, texture, and appearance appear to be related to the
concentration, molecular weight and structure of the polysaccharide. The health benets of b-glucans, such as reducing blood serum
cholesterol and regulating blood glucose levels, are also correlated with the amount and molecular weight of the solubilised b-glucans in
the gastro-intestinal tract.
r 2007 Elsevier Ltd. All rights reserved.
Keywords: Cereal b-glucans; Structure; Molecular weight; Function; Rheology; Gels; Viscoelasticity; Applications; Physiological effects
Contents
1.
2.
3.
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Structural features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Physical properties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.1. Solubilitysolution behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.2. Aggregation phenomenagelation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
102
103
105
105
106
Abbreviations: DP, degree of polymerisation; DP 3, 3-O-b-cellobiosyl-D-glucose; DP 4, 3-O-b-cellotriosyl-D-glucose; DPX5, cellodextrin-like

oligosaccharides with more than three consecutive 4-O-linked glucose residues; DSC, differential scanning calorimetry; FI, uidity index; f, frequency (Hz);
G 0 , storage (or elastic) modulus (Pa); G 00 , loss modulus (Pa); GI, glycemic index; Glcp, D-glucopyranosyl; G 0 max, pseudo-plateau G 0 value (Pa); Gt, gelation
time (h); HDL-cholesterol, high-density-lipoprotein-cholesterol; IE, elasticity increment (h1); IL-1b, interleukin 1b; LDL-cholesterol, low-density-lipoproteincholesterol; Mw, molecular weight; o/w, oil-in-water emulsion; PBGR, peak blood glucose rise; TBG, total b-glucan content; tand, loss tangent ( G 00 /G 0 );
UDP-Glc, uridine diphosphoglucose; g_ , shear rate (s1); g_ 1=2 , shear rate at which Z Z0/2; DG, peak blood glucose level; DH, apparent melting enthalpy (mJ/
mg); Z, apparent viscosity (Pa s); Z*, complex viscosity (Pa s); [Z], intrinsic or limiting viscosity (dl/g); Z0, viscosity at the Newtonian zone (Pa s); (Zsp)0, zero
shear specic viscosity
Corresponding author. Tel.: +30 2310 991716; fax: +30 2310 991797.
E-mail address: athlazar@agro.auth.gr (A. Lazaridou).
0733-5210/$ - see front matter r 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jcs.2007.05.003
ARTICLE IN PRESS
A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118
102
4.
5.
3.2.1. Dynamic rheometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3.2.2. Calorimetry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.2.3. Uniaxial compression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Functional properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.1. Technological aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.2. Physiological responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.2.1. Effect of viscosity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.2.2. Effects of the food matrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.2.3. Fate of b-glucan in the gastro-intestinal tract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4.2.4. Future perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1. Introduction
Over the last two decades the acceptance of b-glucans as
functional, bioactive ingredients has increased the popularity and consumption of cereal-based foods as well as of
many other foods fortied with cell wall-enriched grain
fractions, b-glucan concentrates and isolates. In this
respect, fractions rich in b-glucan have been obtained
from cereal grains by dry milling (pin or roller), sieving,
and air classication processes (Izydorczyk et al., 2003;
Knuckles and Chiu, 1995; Vasanthan and Bhatty, 1995) or
wet milling, sieving, and solvent-extraction protocols using
different solvent systems (Beer et al., 1996; Bhatty, 1995;
Cavallero et al., 2002; Jaskari et al., 1995; Oste Triantafyllou, 2000; Wood et al., 1989). These approaches result in
concentrates or isolates containing approximately 830%
b-glucans for the former and up to 95% for the latter. The
appearance of such ingredients in the market has recently
stimulated the research and development of many novel
food products rich in cereal b-glucans. From a nutritional
and a functional viewpoint, such foods t into the
description of functional foods as they provide some of
the normal quality attributes of a food, such as mouthfeel
and texture, as well as conferring specic health benets.
b-Glucans are major components of starchy endosperm
and aleurone cell walls of commercially important cereals,
such as oat, barley, rye and wheat. The localisation of
b-glucans in cereal grains inuence the isolation and
purication procedures, which aim to produce fractions/
preparations enriched in b-glucans. In oat and barley,
b-glucans are located throughout the starchy endosperm
whereas in wheat the highest concentration is in the
subaleurone layer with little in the rest of the starchy
endosperm (Izydorczyk and Biliaderis, 2000).
b-Glucans from cereals are linear homopolysaccharides
of D-glucopyranosyl residues (Glcp) linked via a mixture of
b-(1-3) and b-(1-4) linkages, with blocks of consecutive
(1-4)-linked residues (i.e. oligomeric cellulose segments)
separated by single (1-3)-linkages (Fig. 1). Although most
of the cellulose segments are trimers and tetramers, longer
cellodextrin units are also present in the polymeric chains
(Dais and Perlin, 1982; Izydorczyk et al., 1998a; Varum
and Smidsrod, 1988; Wood et al., 1991a, 1994a; Wood-
107
108
109
110
110
111
111
112
113
114
114
114
114
ward et al., 1983a, 1988). Cereal b-glucans exhibit

considerable diversity in their structures, including the
ratio of tri- to tetramers, the amount of longer cellulosic
oligomers and the ratio of b-(1-4):b-(1-3) linkages
(Izydorczyk and Biliaderis, 2000). These structural features
appear to be important determinants of their physical
properties, such as water solubility, viscosity, and gelation
properties (Bohm and Kulicke, 1999a, b; Cui et al., 2000;
Doublier and Wood, 1995; Irakli et al., 2004; Izydorczyk
et al., 1998ac; Lazaridou and Biliaderis, 2004; Lazaridou
et al., 2003, 2004; Skendi et al., 2003; Storsley et al., 2003;
Tosh et al., 2004a, b; Vaikousi et al., 2004), as well as of
their physiological action in the gastro-intestinal tract
(Wood, 2002).
The water-soluble bre seems to improve blood glucose
regulation and reduce serum cholesterol levels in diabetic
and hypercholesterolemic subjects, respectively. Such
benecial health effects have been attributed to the
solubility of b-glucans in water and their capacity to form
highly viscous solutions (Kahlon et al., 1993; Wood et al.,
1994b). In 1997, the US Food and Drug Administration
(FDA) approved a health claim for the use of oat-based
foods for lowering the risk of heart disease and passed a
unique ruling that allowed oat bran to be registered as the
rst cholesterol-reducing food at a dosage of 3 g b-glucan
per day, with a recommendation of 0.75 g of b-glucan per
serving (Anonymous, 1997). Recently, a similar health
claim for the barley b-glucan has also been approved
(Anonymous, 2005). However, cereal b-glucans exhibit
only partial solubility in water. Therefore, in addition
to the physiological benets of the soluble dietary bre,
their action also includes the physiological effects associated with the consumption of insoluble bre, such as an
increase of faecal bulk and on ability to relieve constipation. The aforementioned dry milling techniques can provide fractions with b-glucan content up to 30%, including
both soluble and insoluble material. Such fractions are
believed to offer the combined benecial physiological
effects of soluble and insoluble dietary bre when incorporated into food systems. On the other hand, increased
b-glucan solubility is observed for concentrates or isolates
obtained by wet milling processes involving aqueous extractions, hydrolytic enzyme treatments (amylases, proteases),
ARTICLE IN PRESS
.4G14G1
103
Cellotriose unit
3G
4
4
1 G1 G1
Cellotetraose unit
3G
4G
4G14G1
Long cellulosic unit (n3)

3G
4G
1[
4G
1]n
3G
4
1 G1.
Lichenase
4G
4
1 G1
4G
3G
4G
4G
DP 3
DP 4
3G
[4G1]n4G1
1
3G
DP5
Fig. 1. Generalised structure of cereal b-glucans and their debranching with lichenase; dotted arrows indicate the lichenase hydrolysis sites on the
polysaccharide chain. G: b-D-glucopyranosyl unit; DP3: 3-O-b-cellobiosyl-D-glucose; DP4: 3-O-b-cellotriosyl-D-glucose; DPX5: cellodextrin-like
oligosaccharides containing more than three consecutive 4-O-linked glucose residues.
sieving/centrifugation and precipitation of the polysaccharide components with alcohol solution.

The potential use of b-glucans as hydrocolloids in the
food industry is based mainly on their rheological
characteristics, i.e. their gelling capacity and ability to
increase the viscosity of aqueous solutions. Thus, b-glucans
can be utilised as thickening agents to modify the texture
and appearance of food formulations or may be used as
fat mimetics in the development of calorie-reduced foods.
b-Glucan-rich fractions from cereals or puried b-glucans
have in fact been successfully incorporated into products
such as breakfast cereals, pasta, noodles and baked goods
(bread, mufns), as well as dairy and meat products
(Anonymous, 2003; Cavallero et al., 2002; Dexter et al.,
2004; Duss and Nyberg, 2004; Hatcher et al., 2005; Hudson
et al., 1992; Inglett, 1990; Izydorczyk et al., 2005; Knuckles
et al., 1997a; Marconi et al., 2000; Morin et al., 2002;
Newman et al., 1990, 1998; Oste Triantafyllou, 2002;
Volikakis et al., 2004).
The main objective of this review is to address recent
research ndings on structurefunction relationships of
cereal b-glucans in the context of their technological and
physiological functionality. It also considers the prospects
for further research on these topics.
2. Structural features
The structural characteristics of cereal b-glucans of
different botanical origin, including molecular weight,
distribution of cellulose oligomers and ratios of tri- to
tetramers and of b-(1-4) to b-(1-3) linkages in the
polysaccharide chain, are summarised in Table 1.
Despite the structural similarity of b-glucans from
different genera of cereals, as suggested from methylation
analysis and their almost identical NMR spectra, oats,
barley and wheat b-glucans are in fact structurally distinct,
as shown by quantitative HPLC analysis of lichenasereleased oligosaccharides (Cui et al., 2000; Dais and Perlin
1982; Lazaridou et al., 2004; Wood et al., 1991a). The
enzyme lichenase, a (1-3)(1-4)-b-D-glucan-4-glucanohydrolase (EC 3.2.1.73), specically cleaves the (1-4)glycosidic linkages of the three-substituted glucose residues
in b-glucans, yielding oligomers with different degrees of
polymerisation (DP) (Fig. 1). The major hydrolysis
products for the cereal b-glucans are 3-O-b-cellobiosyl-Dglucose (DP 3) and 3-O-b-cellotriosyl-D-glucose (DP 4), but
cellodextrin-like oligosaccharides are also released in
smaller amounts (510%) from the polymer regions
containing more than three consecutive 4-O-linked glucose
residues. The DP of the long cellulose-like fragments has
been found to vary between 5 and 20, with DP5, 6, and 9
being the most abundant (Izydorczyk et al., 1998a, b;
Lazaridou et al., 2004; Wood et al., 1994a). Furthermore,
cellotriosyl and cellotetraosyl segments are distributed in a
rather random pattern in the polymer (Buliga et al., 1986;
Staudte et al., 1983).
The oligosaccharide distribution within the same genera
of cereals is generally similar with major differences only
occurring between b-glucans of different botanical origin
(Cui et al., 2000; Lazaridou et al., 2004; Wood et al.,
1991a). The relative amount of trisaccharide (DP3) in the
b-glucans decreases from wheat (6772%), to barley
(5269%) and oats (5361%), whereas the relative amount
of tetrasaccharide (DP4) follows the opposite trend, i.e.
increases from wheat (2124%), to barley (2533%) and
oats (3441%) (Table 1). However, the combined contents
of tri- and tetrasaccharides are similar among b-glucans
from different cereal genera, resulting in similar total
amount of cellulose-like oligomers with DPX5. The
differences in the proportions of tri- and tetrasaccharides
observed among different b-glucans from various sources
are also reected in the molar ratio of cellotriose to
cellotetraose units (DP3:DP4), following the order of
wheat (3.04.5), barley (1.83.5), rye (1.93.0) and oats
(1.52.3). This ratio is considered to be a ngerprint of
the structure of cereal b-glucans. Literature data indicate
that there also are some differences in the ratio of
DP3:DP4 within the same genera, which could be
ARTICLE IN PRESS
104
Table 1
Molecular-structural features of cereal b-glucans
Source
DP3a
DP4a
DPX5a
Molar ratio
DP3/DP4
(1-4)/(1-3)
Oat
55.058.1
34.236.0
7.78.9
2.12.3
2.32.6
2.4
3603100
57.6
58.3
53.453.8
34.1
33.5
40.441.4
8.2
8.1
1.52.3
1.7
2.2
1.71.8
2.5
2.4
1500
11001500
600840
12002500
1202400
1160
11001600
214257
56.7
55.655.9
34.6
33.634.4
8.7
7.17.5
2.2
1.61.7
2.4
6111700
54.260.9
33.836.7
3.69.7
2.02.3
2.42.8
65250
54.656.8
35.336.3
7.79.2
2.02.1
2.32.6
180850
20602300
1.92.3
5661
2832
613
2.32.9
2.32.6
2.22.6
150290
62.1
29.4
8.4
2.83.4
2.4
17002700
59.264.9
25.330.4
9.410.2
2.63.4
2.4
2.4
80150
13001500
~200600
5702340
56.861.6
63.7
51.861.9
26.132.3
28.5
28.132.1
10.611.2
7.8
6.312.5
1.82.4
3.3
2.32.8
31560
100375
708
66.0
61.564.3
59.464.3
57.762.4
62.063.3
62.069.3
25.7
27.930.1
24.831.0
29.432.9
27.529.2
26.229.1
8.2
7.88.6
8.217.5
7.79.5
8.89.1
4.58.9
3.4
2.73.0
2.53.2
2.32.8
2.83.0
2.83.5
1.92.2
2.22.7
2.12.8
693
1320450
213
250
2.73.0
1.92.3
3.03.8
2.3
1100
21
Wood et al. (1991ac)

Roubroeks et al. (2000b)
Wood et al. (1991a)
72.3
21.0
6.7
4.5
267487
67.1
24.2
8.7
3.7
209
Cui et al. (2000) and Li

et al. (2006)
Lazaridou et al. (2004)
Barley
Rye
Wheat
Molecular
weight (103)
References
Dais and Perlin, (1982)

Doublier and Wood (1995)
and Wood et al. (1991ac)
Autio et al. (1992)
Malkki et al. (1992)
Miller and Fulcher (1995)
Westerlund et al. (1993)
Jaskari et al. (1995)
Beer et al. (1997a, b)
Izydorczyk et al. (1998b)
Zhang et al. (1998)
Cui et al. (2000)
Johansson et al. (2000)
Roubroeks et al. (2000a,
2001)
Wang et al. (2002, 2003)
Colleoni-Sirghie et al.
(2003a)
Lazaridou et al. (2003,
2004)
Skendi et al. (2003)
Aman et al. (2004)
Balance and Manners
(1978)
Dais and Perlin (1982)
Woodward et al. (1983b,
1988)
Wood et al. (1991ac) and
Wood (1994)
Saulnier et al. (1994)
Henriksson et al. (1995)
Beer et al. (1997a)
Gomez et al. (1997a)
Knuckles et al. (1997b)
Izydorczyk et al. (1998a, c)
Morgan and Ofman (1998)
Bohm and Kulicke (1999a)
Cui et al. (2000)
Jiang and Vasanthan
(2000)
Wang et al. (2003)
Wood et al. (2003)
Storsley et al. (2003)
Irakli et al. (2004)
Lazaridou et al. (2004)
Vaikousi et al. (2004)
a
Hydrolysis products of cereal b-glucans by lichenase: DP3 is 3-O-b-cellobiosyl-D-glucose, DP4 is 3-O-b-cellotriosyl-D-glucose and DPX5 is
cellodextrin-like oligosaccharides containing more than three consecutive 4-O-linked glucose residues.
attributed to genotypic and environmental factors (Jiang

and Vasanthan, 2000; Miller et al., 1993; Storsley et al.,
2003; Wood et al., 2003). b-Glucans in waxy barley
varieties appear to have a higher DP3:DP4 ratio compared

to those from non-waxy cultivars. Moreover, the ratios of
tri- to tetrasaccharides in b-glucans from oats and barley
ARTICLE IN PRESS
aleurone tissues are higher than those from starchy

endosperm tissues (Izydorczyk et al., 2003; Wood et al.,
1994a). According to a current model for the biosynthesis
of mixed linkage (1-3),(1-4)-b-glucans proposed by
Buckeridge et al. (2004), the (1-3),(1-4)-b-glucan
synthase makes cellobiosyl and even-numbered cellodextrin units, and a distinct glycosyl tranferase adds a third
glycosyl residue to complete the cellotriosyl and higher
odd-numbered units. Buckeridge et al. (1999, 2001)
determined the b-glucan synthase activity in vitro and
suggested that the cellodextrin unit distribution is altered
drastically depending on the uridine diphosphoglucose
(UDP-Glc) concentration. At suboptimal UDP-Glc concentrations the synthesis of longer cellodextrin units in
b-glucan is favoured, particularly cellotetraosyl units,
whereas as saturation with UDP-Glc is approached,
cellotriose units become the predominant oligomeric
species added along the polysaccharide chain via single
(1-3)-b-linkages.
The calculated ratios of the two types of linkages, (1-4)
to (1-3), in the native cereal b-glucan structures, based on
NMR data and methylation analysis, were found to be
within the range of 1.92.8 (Table 1). Furthermore,
molecular weight values for b-glucans in the range of
653100 103, 312700 103, 211100 103, and 209
487 103 have been reported for oats, barley, rye, and
wheat, respectively. The apparent discrepancies in the
molecular weight estimates of cereal b-glucans may
originate from varietal and environmental (growth) factors, as well as from differences in the methods used for
extraction (solvent and temperature affect the solubilisation) and purication, aggregation phenomena (dependent
on the structural features and solvent quality) and
depolymerisation events (by endogenous or microbial
b-glucanases from contaminating microorganisms) occurring during the extraction step. Also, the reported
molecular weights are dependent on the analytical methodology used for determination of molecular size (including detector and the standards used) (Beer et al., 1996,
1997a, b; Colleoni-Sirghie et al., 2003b; Cui et al., 1999;
Gomez et al., 1997a; Izydorczyk et al., 1998a, c, 2003;
Jaskari et al., 1995; Knuckles et al., 1997b; Malkki et al.,
1992; Morgan and Ofman, 1998; Skendi et al., 2003;
Storsley et al., 2003; Wang et al., 2002; Wood et al., 1989,
1991c; Zhang et al., 1998).
3. Physical properties
3.1. Solubilitysolution behaviour
The chemical features of cereal b-glucans are reected by
their solubility in water and their extended, exible chain
conformation (Woodward et al., 1983b). The cellulose-like
segments of cereal b-glucans might contribute to the
stiffness of the molecules in solution (Varum and
Smidsrod, 1988); b-glucans containing blocks of adjacent
b-(1-4) linkages may exhibit a tendency for interchain
105
aggregation (and hence lower solubility) via strong hydrogen bonds along the cellodextrin portions. The b-(1-3)
linkages break up the regularity of the b-(1-4) linkage
sequence, making the molecule more soluble and exible
(Buliga et al., 1986). It has been suggested that the irregular
spacing of (1-3) linked b-glucosyl residues in the b-glucan
chain is responsible for the non-ordered overall conformation of the polysaccharide and hence the chains are unable
to align closely over extended regions, keeping the
polysaccharide in solution (Woodward et al., 1988). On
the other hand, it has been reported that helical segments
made up of at least three consecutive cellotriosyl residues
may constitute a conformationally stable motif (ordered
domain) in mixed linked (1-3),(1-4) b-glucans (Tvaroska et al., 1983) and that the b-(1-3) linkages could be
involved in the ordered conformation of barley b-glucans
(Morgan et al., 1999). It is possible, therefore, that a higher
content of consecutive cellotriosyl units might impose some
conformational regularity on the b-glucan chain, and
consequently a higher degree of organisation of these
polymers in solution (i.e. lower solubility) (Izydorczyk
et al., 1998c, 2003). In accordance with these two aggregation mechanisms, it has also been suggested that higher
amounts of cellotriosyl fragments and a higher ratio of
(1-4):(1-3) linkages, might explain the solubility differences between cereal b-glucan fractions obtained by using
different aqueous or alkali extraction conditions and/or
different concentrations of ammonium sulphate (Cui et al.,
2000; Izydorczyk et al., 1998ac; Storsley et al., 2003;
Woodward et al., 1988).
Dynamic and steady shear rheological tests of freshly
prepared (or heat-treated before measurement) solutions of
cereal b-glucans reveal a behaviour typical of noninteracting disordered polysaccharides with chain entanglements in the concentrated state. Thus, for aqueous
dispersions of b-glucan a Newtonian region at low shear
rates and a shear-thinning ow at high shear rates are
observed (Fig. 2a). Moreover, under dynamic rheological
measurements the loss modulus, G00 , is larger than the
storage (or elastic) modulus, G0 at lower frequencies, while
the behaviour approaches that of solid-like materials at
higher frequencies, with G0 being greater than G00 . As
expected, with increasing molecular weight (Fig. 2a) and/or
concentration of polysaccharide, there is an increase in
viscosity as well as in the shear thinning and viscoelastic
properties of aqueous dispersions of b-glucans (Irakli et al.,
2004; Lazaridou et al., 2003, 2004; Skendi et al., 2003;
Vaikousi et al., 2004).
The reported intrinsic or limiting viscosity ([Z]) values for
cereal b-glucans vary between 0.28 and 9.6 dl/g, depending
largely on the molecular weights of the isolated polysaccharides. It has been suggested that b-glucans have an
extended random coil conformation. Consequently, the
double logarithmic plots of zero shear specic viscosity,
(Zsp)0, vs. c[Z] for several b-glucan preparations differing in
ne structure and molecular size superimpose closely, often
falling into three linear parts (Bohm and Kulicke 1999a;
ARTICLE IN PRESS
106
-3
Mw x 10
250
210
180
140
110
70
10
0.1
/0
(Pa s)
Mw x 10-3 DP3/DP4
0.01
0.1
0.001
0.01
0.01
(1/s)
100
200
200
200
100
100
0.0001
0.001 0.01 0.1
10000
2.1
3.0
3.7
2.1
2.8
1
10
1/2
100 100010000
Fig. 2. Molecular weight (70250 103 Mw) effects on apparent viscosity (Z) of barley b-glucan aqueous dispersions (3% w/v, 20 1C and DP3:DP4 ratio
2.8) (adapted from Vaikousi et al., 2004) (a) and generalised shear rate dependence of viscosity for cereal b-glucans (8% w/v, 20 1C); the solid line
represents the t of the experimental data to Eq. (1) (adapted from Lazaridou et al., 2004) (b).
Burkus and Temelli 2003; Colleoni-Sirghie et al., 2003b;

Doublier and Wood 1995; Grimm et al., 1995; Gomez
et al., 1997b; Irakli et al., 2004; Izydorczyk et al., 1998a, b;
Lazaridou et al., 2003, 2004; Roubroeks et al., 2000a, b;
Skendi et al., 2003; Vaikousi et al., 2004; Varum and
Smidsrod 1988; Wang et al., 2003; Woodward et al., 1983a,
1988).
The ow curves of aqueous solutions of b-glucans have
been described by the power law model and are largely
depended on molecular size, concentration and temperature. Moreover, the effects of temperature on the viscosity
of b-glucan solutions have been modelled by the Arrhenius
equation (Autio et al., 1988; Colleoni-Sirghie et al., 2003b;
Papageorgiou et al., 2005; Wikstrom et al., 1994). The
shear-thinning curves for aqueous solutions of b-glucans
were found to follow a common master curve using a
transformation of viscosity values vs. shear rate (Fig. 2b).
This general form of shear-thinning behaviour is typical of
disordered polysaccharides irrespective of primary structure, molecular weight, temperature, solvent environment,
and concentration, and can be matched with good
precision by the simple relationship (Morris, 1990):
Z Z0 =1 _g=_g1=2 0:76 ,
(1)
where Z is viscosity, Z0 is viscosity at the Newtonian zone, g_

is shear rate, and g_ 1=2 is shear rate at which Z Z0/2.
Indeed, the experimental data from the generalised shearthinning prole of cereal b-glucan solutions at certain
concentrations (8, 10% w/v) and temperature (20 1C) tted
well (r2 0.99) to this equation regardless of differences in
the molecular weight and ne structure of the b-glucan
isolates (Lazaridou et al., 2004; Vaikousi et al., 2004). The
application of the CoxMerz rule (i.e. the relationship
between rheological responses under destructive and nondestructive deformation) has also been tested for cereal
b-glucan preparations, differing in molecular weight and
in the concentration of dissolved polysaccharide. This rule

is generally observed for random coil polymer chains
interacting solely by physical entanglements and no large
departures from the empirical CoxMerz correlation were
found for cereal b-glucan samples, except in the case of
high polysaccharide concentrations and samples of high
molecular weight (Autio, 1988; Bohm and Kulicke 1999a;
Lazaridou et al., 2003, 2004; Skendi et al., 2003).
3.2. Aggregation phenomenagelation
Although fresh solutions of cereal b-glucans fall
rheologically into the category of random coil-type
polysaccharides, their rheological properties may change
depending on their molecular characteristics (size, structure), storage time (i.e. waiting time before analysis) and
thermal history (Bohm and Kulicke 1999a, Gomez et al.,
1997c; Lazaridou et al., 2003, 2004; Tosh et al., 2003;
Vaikousi et al., 2004). Cereal b-glucans with certain
structural features may display time-dependent rheological
behaviour, often revealed in thixotropic loop experiments,
implying a time-dependent formation of intermolecular
networks. Thus, with increasing storage time solutions of
b-glucans show unusual shear-thinning ow behaviour at
low shear rates. This behaviour becomes more pronounced
with increasing polysaccharide concentration and storage
time prior to the rheological testing, as well for b-glucans
with low molecular weight and high proportions of
cellotriosyl units in the polymeric chains (Lazaridou
et al., 2003, 2004; Vaikousi et al., 2004).
In addition to exhibiting increased solution viscosity on
storage, b-glucans have also been shown to gel under
certain conditions. Thus, cereal b-glucan hydrogels with
different molecular characteristics and properties have
been obtained under isothermal (545 1C, 412% w/v
polymer concentration) conditions (Bohm and Kulicke
1999b; Irakli et al., 2004; Lazaridou et al., 2003, 2004; Li
ARTICLE IN PRESS
et al., 2006; Papageorgiou et al., 2005; Skendi et al., 2003;

Tosh et al., 2003, 2004a; Vaikousi et al., 2004), as well as
after repeated freezing and thawing cycles of relatively
dilute (14% w/v) polysaccharide solutions due to aggregation of the polymeric chains upon freeze-concentration
(Lazaridou and Biliaderis 2004; Vaikousi and Biliaderis,
2005). The later process is called cryogelation or cryostructurisation and the gels formed cryogels. Cereal
b-glucan hydrogels formed at temperatures above 0 1C
and cryogels belong to the category of physically crosslinked gels whose three-dimensional structure is stabilised
mainly by multiple inter- and intrachain hydrogen bonds in
the junction zones of the polymeric network. The structure
and properties of b-glucan hydrogels have been explored
by small strain dynamic rheometry, differential scanning
calorimetry (DSC) and large deformation mechanical tests.
3.2.1. Dynamic rheometry
The gelation capacity of aqueous dispersions of cereal
b-glucans differing in molecular size and ne structure has
been monitored isothermally by dynamic rheometry at
different temperatures above 0 1C and polymer concentrations (Irakli et al., 2004; Lazaridou et al., 2003, 2004; Li
et al., 2006; Skendi et al., 2003; Vaikousi et al., 2004). After
an induction period, G0 and G00 increase with time and the
aqueous dispersions begin to adopt gel-like properties
(G0 4G00 ), as shown in Fig. 3. At the end of the gel curing
process, the behaviour becomes typical of an elastic gel
network and the G0 attains a pseudo-plateau value G0 max.
The time where G0 crosses G00 is often considered as the
gelation time (Gt) and the maximum slope of the G0 curve is
reported as an index of the gelation rate; the latter, known
as elasticity increment, IE, can be calculated as
IE (dlog G0 /dt)max (Bohm and Kulicke 1999b). The Gt
1000
0.1
0.01
0.1
10
f (Hz)
0.1
100
Gmax
1000
G
100
G
1000
1000
G
10
IE = (dlogG/dt)max
100
40
60
80
Time (h)
10
0.1
20
10
Gt
100
(Pa s)
G, G (Pa)
10
(Pa s)
G, G (Pa)
G, G (Pa)
decreases and the IE values increase with decreasing

molecular weight in cereal b-glucans with similar distributions of cellulose-like fragments (Fig. 4a), possibly due to
the higher mobility of the shorter chains that enhances
diffusion and lateral interchain associations (Bohm and
Kulicke 1999b; Doublier and Wood 1995; Irakli et al.,
2004; Lazaridou et al., 2003, 2004; Li et al., 2006; Skendi
et al., 2003; Tosh et al., 2004b; Vaikousi et al., 2004).
Among cereal b-glucans of equivalent molecular weight,
the gelation time decreases and the gelation rate increases
in the order of oats, barley, and wheat b-glucans, reecting
the order of the molar ratio of DP3:DP4 units (Fig. 4b)
(Bohm and Kulicke 1999b; Cui et al., 2000; Lazaridou et
al., 2004; Tosh et al., 2004a). Two different gelation models
have been proposed in the literature for mixed-linkage (1-3),
(1-4) b-glucans. One involves the side-by-side interactions
of cellulose-like segments of more than three contiguous
b-(1-4)-linked glucosyl units (Fincher and Stone 1986)
and the other the association of chain segments with
consecutive cellotriosyl units linked via b-(1-3) bonds
(Bohm and Kulicke 1999b). Overall, the observed differences in gelation propensity seem to be attributed mostly to
the differences in the DP3:DP4 ratio between cereal
b-glucans; the likelihood of ordered repeating segments
of the cellotriosyl units being increased with an increase in
the DP3:DP4 ratio. Furthermore, a decrease in molecular
weight and an increase in the DP3:DP4 ratio in the
b-glucan chains resulted in gels of increased G0 max and
decreased values of loss tangent, tan d ( G00 /G0 ), (Lazaridou et al., 2004; Tosh et al., 2004a). The gelation rate also
increases with increasing concentration and increasing gel
curing temperature, reaching a maximum at 2535 1C.
At higher temperatures the IE values decrease (Lazaridou
et al., 2003, 2004; Skendi et al., 2003; Vaikousi et al., 2004).
10000
10
100
107
100 120
140
1
0.01
1
0.1
1
f (Hz)
10
100
Fig. 3. Solgel phase transition of an aqueous b-glucan dispersion (10% w/v) as probed by dynamic rheological testing (gel curing at 25 1C, strain 0.1%,
frequency 1.0 Hz); the respective mechanical spectra before (left) and after (right) gelation are also shown.
ARTICLE IN PRESS
108
60
Gt
IE
50
180
2.5
160
2
0.2
140
IE (h-1)
30
Gt (h)
1.5
0.15
100
80
0.1
Gt (h)
120
40
IE (h-1)
0.25
Gt
IE
60
20
40
0.5
10
0.05
20
0
0
0
50 100 150 200 250 300
0
2
2.5
Mw x10-3
3
3.5
DP3/DP4
Fig. 4. Molecular weight (a) and structure (b) dependence of elasticity increment (IE) and gelation time (Gt) for cereal b-glucans cured at 25 1C and 8% w/v
concentration (frequency 1 Hz, strain 0.1%); the b-glucan preparations of (a) have a common DP3:DP4 ratio, 2.8, and those of (b) have a common
molecular weight, 210 103 (adapted from Lazaridou et al., 2004 and Vaikousi et al., 2004).
0.5
Mw x 10-3
0.45
0.4
0.35
DP3/DP4
65
2.1
110
2.1
140
200
2.1
2.1
200
3.0
200
3.7
0.3
tan
Furthermore, the dependence of the storage modulus

(apparent G0 max value) on b-glucan concentration follows
power law relationships; G0 varying as C7.27.5 (Lazaridou
et al., 2003). For cereal b-glucan cryogels, the G0 values
obtained from their mechanical spectra at 5 1C increase and
tan d decrease with increasing initial polysaccharide concentration, number of freezethaw cycles and trisaccharide
units (DP3), as well as with decreasing molecular weight of
the polysaccharide (Fig. 5) (Lazaridou and Biliaderis,
2004).
Food formulation also has an impact on the gelation
ability and the rheological properties of cereal b-glucan
gels (Irakli et al., 2004; Lazaridou and Biliaderis, 2007;
Lazaridou et al., 2007; Vaikousi and Biliaderis, 2005). The
incorporation of various sugars (glucose, fructose, sucrose,
xylose, and ribose) at a concentration of 30% (w/v) to
aqueous dispersions of barley b-glucan (6% w/v) led to an
increase of the gelation time upon curing at room
temperature (Irakli et al., 2004). Similarly, the addition of
several sugars (sucrose, glucose, fructose, and xylose) at
a concentration range of 530% (w/w) to aqueous dispersions of b-glucan (3% w/w) without or with skimmed
milk (12% w/w), submitted to repeated cycles of freezingthawing, induced a signicant delay in the transition
to a gel state. The resultant cryogels were weaker
(Lazaridou and Biliaderis, 2007; Lazaridou et al., 2007;
Vaikousi and Biliaderis, 2005) and the inhibitory effects
of xylose and fructose were stronger than those of glucose
and sucrose. On the other hand, sorbitol appears to
promote network formation (Lazaridou and Biliaderis,
2007; Lazaridou et al., 2007). However, whereas the
effects of polyols were signicant when added to preparations of high molecular weight b-glucans (210 103), they
generally decreased with decreasing molecular weight and
seemed to be independent on the type of polyol for
low molecular weight b-glucan preparations (e.g. 140 and
70 103).
0.25
0.2
0.15
0.1
0.05
0
0
9
6
Number of cycles
12
Fig. 5. Cereal b-glucan molecular weight and structure effects on tan d

recorded at 5 1C, 1.06 Hz, and 0.1% strain of 3% (w/v) cryogels obtained
after repeated freezethaw cycles (adapted from Lazaridou and Biliaderis,
2004).
3.2.2. Calorimetry
The DSC thermal scans of cereal b-glucan hydrogels
formed by curing at temperatures above 0 1C (Fig. 6a) and
by freezethaw cycling exhibited rather broad endothermic
gel-sol transitions at 5580 1C (Lazaridou and Biliaderis,
2004; Lazaridou et al., 2003, 2004; Skendi et al., 2003;
Vaikousi et al., 2004). Similar to the rheological responses,
the DSC kinetic data showed that the rate of endotherm
development (short range molecular order) during gel
curing at room temperature (Fig. 6b) (Lazaridou et al.,
ARTICLE IN PRESS
Mwx10-3 DP3/DP4
curing: 21h
35
Mw x 10-3 DP3/DP4
35
2.3
65
2.3
200
2.1
200
3.0
200
3.7
2.3
5
H: 5.1 mJ/mg
4
65
curing: 47h
2.3
H: 2.9 mJ/mg
0.2 mWatts
curing: 72h
110
2.3
110
2.8
H (mJ/mg)
Endothermic Heat Flow
109
H: 2.1 mJ/mg
curing: 40h
1
H: 4.1 mJ/mg
0
30
40
50
60
70
80
Temperature (C)
90
100
20
40
60
Time (h)
80
100
120
Fig. 6. Cereal b-glucan molecular weight and structure effects on differential scanning calorimetry (DSC) thermal curves of gels cured for specied time
periods (a), and on apparent melting enthalpy (DH) kinetic data of gel structures (b); heating rate 5 1C/min and curing at 25 1C with 10% w/v
polysaccharide concentration (adapted from Lazaridou et al., 2003, 2004).
2003, 2004), as well as the apparent melting enthalpy values

(plateau DH) of the gels increase with decreasing molecular
size and with increasing DP3:DP4 ratio (Fig. 6) (Lazaridou
et al., 2003, 2004; Li et al., 2006; Vaikousi et al., 2004). The
DH values of the cryogels also increase with increasing
DP3:DP4 ratio and with decreasing molecular weight of
cereal b-glucans (Lazaridou and Biliaderis, 2004). Furthermore, the melting temperatures of the gel networks studied,
as measured by dynamic rheometry and DSC, increased
with the molecular weight and the amount of cellotriosyl
units in the polysaccharide chains (Lazaridou and Biliaderis, 2004; Lazaridou et al., 2003, 2004; Li et al., 2006;
Vaikousi et al., 2004). It appears that although a slower
gelation process is observed for high molecular weight
b-glucans, the gel network structure consists of structural
elements (microaggregates) with better organisation and/
or involves interchain associations over longer chain
segments.
It is also notable that cryogels obtained after repeated
freezethaw cycles, at low initial polysaccharide concentrations (3% w/v) (Lazaridou and Biliaderis, 2004), are
analogous with the gel networks prepared at room
temperature and using much higher polysaccharide concentrations (10% w/v) (Lazaridou et al., 2004). In fact, the
DSC thermal scans of these two systems show that the
apparent melting enthalpy (DH) values are similar for both
types of gels whereas comparison of the broadness of the
endotherm peak reveals that cryogelation generally yields
a more cooperative gel-sol transition. Similarly, after
curing 3% (w/v) b-glucan dispersions at 5 1C for 135 days,

which is long enough for G0 to reach a pseudo-plateau
value, the strength (G0 value) of the resultant gel is lower
than that of cryogels obtained from dispersions at
equivalent initial polysaccharide concentration after 14
freezethaw cycles (Lazaridou et al., 2007). The reinforcement of b-glucan network structures induced by cryostructurisation is attributed to the increased concentration
(cryoconcentration) of the polysaccharide in the unfrozen
bulk phase resulting in the promotion of associative
interactions among the polysaccharide chains (physical
cross-linking).
3.2.3. Uniaxial compression
Variation in the mechanical properties of cereal b-glucan
gels has been also revealed by large deformation compression tests. For gel samples cured at room temperature, an
increase in strength and a decrease in brittleness were
found with increasing concentration, molecular weight and
DP3:DP4 ratio of the polysaccharide (Lazaridou et al.,
2003, 2004; Vaikousi et al., 2004). Instead, cereal b-glucans
with high molecular weight and low amounts of DP3 units
formed strong cryogels when tested under large deformation protocols; these ndings were attributed to differences
in the nature of the network microstructure between
samples formed under different gel curing conditions
(Lazaridou and Biliaderis, 2004). Microscopic images of
cereal b-glucan gels aged for seven days at 5 1C revealed
that the microstructure was not homogenous and there
ARTICLE IN PRESS
110
seemed to be a coarsening of the gel network structure as

the b-(1-3)-linked cellotriosyl unit content increased
(Tosh et al., 2004a).
4. Functional properties
4.1. Technological aspects
The incorporation of fractions enriched with oats and
barley b-glucans into cereal-based foods has been widely
studied. The addition of b-glucan concentrates at high
levels (1030%) in yeast-leavened breads leads to high
water retention, as well as to darker coloured end-products
and undesirable effects on crumb texture and loaf volume
(Bhatty, 1986; Cavallero et al., 2002; Dexter et al., 2004;
Izydorczyk and Dexter, 2004; Knuckles et al., 1997a;
Krishnan et al., 1987; Newman et al., 1998). However,
b-glucan rich fractions from oats and barley may result in
increased loaf volumes when used at certain concentrations
in wheat our, probably by increasing the viscosity of the
dough (Delcour et al., 1991). It is considered that b-glucans
may play a role in improvement of bread crumb structure
by stabilising air cells in the dough and preventing their
coalescence (Wang et al., 1998). Relationships have also
been established between b-glucan concentration and the
pasting characteristics of oat ours. The increase in the
pasting peak viscosity which occurred with increasing
b-glucan content of oat cultivars was explained by an
increase in the water binding capacity of the ours
(Colleoni-Sirghie et al., 2004; Zhou et al., 2000). In
addition to the polysaccharide concentration, the molecular weight (Mw) also seems to affect the breadmaking
performance of b-glucans, as shown in a recent investigation (Skendi et al., 2006) in which two isolates of barley
b-glucan with different molecular weights were incorporated into breads made from two types of wheat our
differing in their breadmaking quality. The addition of
b-glucans with high molecular weight resulted in higher
loaf volumes than the incorporation of those with low
molecular weight. The highest loaf volume was obtained
for the poor breadmaking quality our with 0.6% w/w
b-glucan, with the incorporation of the high molecular
weight preparation giving quality exceeding that of the
good breadmaking quality wheat cultivar. Furthermore,
incorporation of b-glucans up to 5% into baked goods,
such as mufns, did not seem to affect the overall
acceptability of the products (Hudson et al., 1992; Newman et al., 1990, 1998). The addition of barley fractions
enriched in b-glucans in semolina or wheat ours at up to
20%, resulted in pasta or noodles with acceptable sensory
properties and cooking quality despite their decreased
brightness and yellowness and the increased redness and
speckiness of the product (Dexter et al., 2004; Hatcher
et al., 2005; Izydorczyk et al., 2005; Knuckles et al., 1997a;
Marconi et al., 2000).
The aforementioned development of weak gel network
structures by cereal b-glucans with certain structural
characteristics and under certain conditions is a desirable

attribute in water-continuous low-fat spreads. In this
context, cereal b-glucans exhibit the potential to be used
as fat replacers due to their high viscosity and waterbinding and their foaming and emulsion stabilising capabilities (Burkus and Temelli, 2000; Kontogiorgos et al.,
2004). Kontogiorgos et al. (2004) investigated the effects of
pure barley and oat b-glucans on the rheological and
creaming behaviour of concentrated egg-yolk-stabilised
model emulsions (salad dressing model). The high molecular weight b-glucans (apparent Mw110 103) stabilised
the oil-in-water (o/w) emulsions, mainly by increasing the
viscosity of the continuous phase, while the low molecular
weight b-glucans (apparent peak Mw40 103) inuenced
emulsion stability through network formation in the
continuous phase. Moreover, it was found that incorporation of barley b-glucan gum (76.2% purity) into reducedfat breakfast sausages at a level that provides 0.30.7%
b-glucan in the product gave improved water binding,
without having any signicant effects on product texture or
avor at a level of 0.3% (Morin et al., 2002). A commercial
concentrate of oat b-glucans (22.2% b-glucan content) has
been also incorporated into low-fat white-brined cheese
from bovine milk (70% fat reduction) at two levels, 0.7%
and 1.4% (w/w), resulting in a product with increased yield,
greater proteolysis and higher levels of short chain fatty
acids (lactic, acetic, and butyric) as well as with improved
texture compared to its low-fat (b-glucan-free) counterpart. However, the colour, avour and overall impression
scores were signicantly inferior to those of a typical whitebrined cheese product, particularly for the cheese product made with the high level of b-glucan (Volikakis
et al., 2004). Recently, non-dairy ready-to-use, lactose-free,
milk substitutes (oat milk beverage, yogurt, ice cream,
oat-based cream, whipped cream, and buttermilk) have
been introduced into the market. These products contain
cereal b-glucans as bioactive ingredients as well as acting
as stabilisers and texturisers (Anonymous, 2003; Oste
Triantafyllou, 2002).
The molecular weight of b-glucans does not only affect the
viscosity of foods but also has a signicant impact on the
perceived (sensory) thickness of products as shown for
beverage and soup food prototypes containing 0.252%
(w/w) b-glucans (Lyly et al., 2003, 2004). These appear to be
signicant correlations between textural characteristics and
measured viscosity as well as a moderately strong relationship
between avor attributes and the viscosity of soups. From a
technological point of view, b-glucans may be useful as
alternative thickening agents in soups. It may also be possible
to prepare beverages and soups containing a physiologically
effective amount of b-glucan from low molecular weight
b-glucan preparations, as they are easier to process than their
high molecular weight counterparts. However, the relationship between the physiological functionality and molecular
weight of b-glucans has to be borne in mind.
The potential use of b-glucans as biodegradable edible
food packaging material has recently been suggested
ARTICLE IN PRESS
(Skendi et al., 2003). The mechanical properties of cast

b-glucan lms were affected by the amount of plasticiser
(water, polyol) used and the molecular weight of the
polysaccharide. Water as well as sorbitol, added as a coplasticiser at a 15% (w/w) level, improved the extensibility,
but decreased the mechanical strength of the b-glucan
lms. Furthermore, high molecular weight b-glucans gave
stronger and more extensible lms than low molecular
weight b-glucan preparations.
4.2. Physiological responses
4.2.1. Effect of viscosity
The physiological functions of dietary bre are often
attributed to their physico-chemical properties: water
holding capacity, swelling, diffusion-suppressing ability
(through viscosity enhancement and gel-formation), binding properties, and susceptibility or resistance to bacterial
degradation and fermentation (Dikeman and Fahey, 2006;
Schneeman, 2001). The mechanisms by which a soluble
bre, such as b-glucan, exerts hypocholesterolemic and
hypoglycemic effects are still debated but the most
common hypothesis is based on increased lumen viscosity
(Battilana et al., 2001; Bell et al., 1999; Bourdon et al.,
1999; Dikeman and Fahey, 2006; Wood, 2002). It has been
suggested that cereal b-glucans decrease the absorption and
reabsorption of cholesterol, bile acids, and their metabolites by increasing the viscosity of the gastro-intestinal tract
contents as well as delaying gastric emptying and the
intestinal absorption of nutrients, such as digestible
carbohydrates, and thereby reducing post-prandial hyperglycemia and insulin secretion. The latter have health
benets for those with type-2 diabetes, and are also
associated with reduced risk of developing the disease
and insulin insensitivity. Carbohydrate and lipid metabolism are closely interrelated and insulin has also been
reported to increase hepatic cholesterol synthesis. Therefore, if bre decreases carbohydrate absorption and insulin
secretion, it may also contribute indirectly to the hypocholesterolemic effects (Bell et al., 1999; Bourdon et al., 1999;
Dikeman and Fahey, 2006; Wood, 2002). Furthermore, the
increased viscosity caused by soluble dietary bre inuences fat emulsication by increasing emulsion droplet size
which may impair fat absorption (Pasquier et al. 1996).
However, Battilana et al. (2001) found that the administration of frequent meals with or without b-glucans
resulted in similar carbohydrate and lipid metabolism,
whereas ingestion of a single meal containing b-glucan
lowered post-prandial glucose concentrations. This might
suggest that the benecial action of b-glucans is mainly due
to delayed and reduced carbohydrate absorption from the
gut and does not result from the effects of metabolites
produced by fermentation of b-glucans in the colon. It is
also likely that b-glucans do not only decrease the postprandial glucose response due to high viscosity in the
gastro-intestinal tract, but also reduce starch digestion by
a-amylase. Symons and Brennan (2004) found that
111
inclusion of 5% of a b-glucan-rich fraction in bread

resulted in a signicant decrease in the release rate of
reducing sugars following the in vitro digestion of bread
with pepsin and a-amylase. Similarly, Izydorczyk et al.
(2005) showed that replacement of 25% wheat our in
noodles with a bre-rich fraction from barley signicantly
decreased the in vitro release of glucose compared to 100%
wheat noodles. In both cases, it was speculated that
b-glucans decrease the accessibility of starch-degrading
enzymes to their starch substrate by forming a gel matrix
or by limiting the water available for starch hydration and
thus, for starch gelatinisation.
A study with ileostomy subjects showed that bile acid
excretion increased by a mean of 450% in a diet with oat
bran bread, compared with a similar diet in which 70% of
the b-glucan was degraded by b-glucanase. This indicates
that the high molecular weight b-glucan mediates increased
excretion of bile acids: whole micelles or bile acids may
become entrapped or encapsulated in the highly viscous bglucan matrix, and then excreted from the small bowel or
the viscous environment to reduce the formation of mixed
micelles in the small intestine and/or decrease the
reabsorption of bile acids in the terminal ileum (Lia
et al., 1995). Furthermore, a dependence of the hypocholesterolemic action on extractability, viscosity, pseudoplastic ow behaviour, hydrodynamic properties, and
molecular weight of b-glucans was found in rats fed on
diets based on various oat bran concentrates, supporting a
mechanism based on high lumen viscosity (Malkki et al.,
1992). A difference in the mechanism of physiological
effects has also been observed between cereal b-glucans
differing in their molecular features. Wilson et al. (2004)
demonstrated that the cholesterol-lowering activity of
barley b-glucan in hamsters occured with both low
(175 kDa) and high molecular weight (1000 kDa) preparations. However, although cholesterol accumulation did not
differ between the molecular weight preparations, the
esterication of cholesterol was affected. Thus, a signicantly lower accumulation of cholesterol ester was observed in the hamsters fed the lower molecular weight
barley b-glucan: tissue cholesterol ester concentration is
viewed as the hallmark in fatty streak formation associated
with atherogenic progression. Several other animal studies
of diets supplemented with cereal bre or cereal fractions
showed that the content of b-glucans and particularly that
of soluble b-glucans, the molecular weight of the b-glucans
and endogenous b-glucanase activity, all of which affect
extract viscosity, appeared to be strong predictors of the
serum cholesterol response (Kahlon et al., 1993; Newman
and Newman, 1991; Newman et al., 1992). In a metaanalysis report, Brown et al. (1999) identied 25 clinical
studies involving a total of 1600 healthy, hyperlipidemic
and diabetic individuals in the age range of 2661. Daily
doses of between 2 and 10 g of oat soluble bre gave
signicant mean reductions in the total (0.04 mmol/L/g of
bre) and low-density-lipoprotein (LDL, 0.037 mmol/L/g
of bre) cholesterol levels. It is generally considered that
ARTICLE IN PRESS
112
the cholesterol-lowering effect is larger in subjects with

higher cholesterol levels, whereas the doseresponse effect
is not always clear. Overall, the ndings from many human
studies reveal a substantial variation in the responses
between the different trials and types of subjects studied. In
addition to variation in the design (intervention) of the
clinical trial (i.e. number and type of subjects involved,
daily intake and number of servings, length of treatment,
etc.), variation is also observed in the physiological
responses of individual. Also, in some cases, inconsistent
results may be related to factors that inuence the viscosity
development by b-glucans, which is in turn determined by
the molecular weight and extent of solubilisation of the
polysaccharide from the food matrix.
According to Anderson (1990) and Malkki and Virtanen
(2001), soluble bre can affect satiety and promote weight
loss for a number of reasons including a slower rate of meal
intake, a delay in gastric emptying, moderate responses of
plasma glucose levels and insulin secretion by the pancreas
(insulin stimulates hunger), elevation of cholecystokinin
(a gut hormone correlated with prolonged satiety) and
production of gas and short-chain fatty acids by fermentation of the bre in the colon.
It has recently been shown that a gelling b-glucan from
barley (Glucagel) can cause a number of effects on the
mammalian immune system, including upregulation of the
release of the cytokine interleukin 1b (IL-1b) (indicating
enhanced monocyte differentiation), changes to wound
healing processes and enhancement of gut mucin secretion
(Porter et al., 2006). Furthermore, the in vitro secretion of
IL-1b, which is one of the early responses of the immune
system to infection, increased with increasing dose and
molecular weight of the b-glucan.
4.2.2. Effects of the food matrix
The viscosity effects of b-glucans are often modulated by
formulation and food processing and storage protocols.
The doseresponse of oat gum on plasma glucose and
insulin levels of healthy humans consuming oat gum in a
drink (in the range of 1.814.5 g after an oral glucose load
of 50 g) has been studied (Wood et al., 1994b). Increasing
the dose of oat gum successively reduced the plasma
glucose and insulin responses relative to a control drink
without gum, reaching a plateau doseresponse at an
intake of 6 g of b-glucan (the maximum effective dose). In
a recent study by Cavallero et al. (2002), a linear decrease
in the glycemic index of non-diabetic humans was found
with increasing consumption of barley b-glucan in bread,
the GI (glycemic index) being related to the total b-glucan
content (TBG) by the equation:
GI 91:27 3:68TBG r2 0:96.
(2)
It should also be noted that the doseresponse for a solid

meal might differ from that for a liquid (Wood et al.,
1994b). The food matrix or food processing or both could
have adverse effects on the hypocholesterolemic properties
of b-glucan. A clinical study (Kerckhoffs et al., 2003) with
mildly hypercholesterolemic subjects showed that consumption of an b-glucan-enriched preparation from oat
bran with orange juice was more effective in lowering total
and LDL-cholesterol concentrations and the ratio of total
to high-density-lipoprotein (HDL)-cholesterol than when
the same preparation was administered in bread and
cookies. Some studies have failed to show a physiological
action of b-glucan preparations despite the daily b-glucan
intake being much higher than the amount recommended
by the FDA. This fact has been attributed to the reduction
of polysaccharide molecular weight occurring during the
isolation stage and/or as a result of food processing (Keogh
et al., 2003). Commercial oat foods including porridge
made of rolled oats, and extruded and breakfast cereal
products all contain high molecular weight b-glucans
(6002930 103), crisp bread (950 103) and yogurt-like
products (830 103) contain moderately degraded
b-glucans while bread loaves, fried pancakes, pancake
batter, and fermented oat soup contained extensively
degraded b-glucans (190630 103) (Aman et al., 2004;
Wood et al., 1991c). During baking of yeast-leavened oatand barley-containing breads, the b-glucan is partially
degraded (from 12002300 103 to 2401920 103), probably due to the action of b-glucanases in the our mixtures.
This degradation is reduced with decreasing mixing and
fermentation time and with increasing particle size of the
bran (Aman et al., 2004; Andersson et al., 2004; Sundberg
et al., 1996; Trogh et al., 2004). Furthermore, during the
typical pasteurisation conditions used for an acidic liquid
product such as fruit and tomato juice, a reduction of
viscosity resulting from the inclusion of barley b-glucan
isolates is observed, probably as a consequence of acid
hydrolysis (Vaikousi and Biliaderis, 2005). The effects of
acid hydrolysis were dependent on pH, temperature and
time, and were more pronounced for a high molecular
weight isolate (molecular weight 250 103) than for a low
molecular weight isolate (140 103), indicating that
differences in the ow behaviour of liquid products
containing b-glucans of different molecular size may occur
under different processing and formulation protocols.
Furthermore, food processing and storage can alter the
solubility and thereby, the availability of b-glucans (Beer
et al., 1997b; Brummer et al., 2006). Food processing can
increase the physiological activity of b-glucans by increasing availability (cooking, extrusion) and although the
molecular size of the polymer may be partly reduced (due
to enzymic hydrolysis, milling, stirring, and pumping)
(Izydorczyk et al., 2000; Robertson et al., 1997), the
b-glucan can be still effective in improving plasma
cholesterol, glucose and insulin responses (Sundberg
et al., 1995; Van Der Sluijs et al., 1999; Yokoyama et al.,
1998). Animal studies have shown that the molecular
weight of b-glucans can be reduced by an order of
magnitude (e.g. from 1000 to 200 kDa) and the preparations are still able to reduce plasma cholesterol levels
(Yokoyama et al., 2002; Wilson et al., 2004). However, a
recent clinical study showed that there were no appreciable
ARTICLE IN PRESS
113
effects on the concentrations of blood lipids, insulin or

glucose in moderate hypercholesterolemic humans on consumption of yeast-leavened oat breads with high (797 kDa)
or low (217 kDa) molecular weight b-glucans (Frank et al.,
2004). It is probable that these responses are due to the
solid nature of the food matrix which can inuence the
solubility and extractability of the added polysaccharide.
Wood (2002) has suggested that the net result of food
processing might be to improve the bioavailability or
bioactivity of the b-glucan, and that this is related to the
product of (C Mw); C being the concentration and Mw
the molecular weight. Human studies have shown inverse
linear relationships between log(viscosity) and post-prandial glucose and insulin responses after consuming glucose
solutions containing oat gum in the dose range of 1.3
10.5 g pure b-glucan (Wood et al., 1994b). The viscosity of
aqueous oat gum solutions which improved post-prandial
glucose and insulin responses when consumed by healthy
humans was within the range of 208000 mPa s (at 30 s1),
with little additional effect being achieved above 5000 mPa s.
Regression analysis of their experimental data suggested
that even at viscosities as low as 10 mPa s, an average
1213% reduction in peak plasma glucose could be
observed. In addition, a relationship between the change
in peak blood glucose levels (DG), the polysaccharide
concentration (c) and the weight average molecular weight
(Mw) of the b-glucan has been found (Wood et al., 2000):
attenuating the glycemic response; this was achieved in

mufns subjected to freezethaw cycles to modify the
solubility of the polysaccharide. The freezethaw treatment
reduced the solubility from 30% to 40% in fresh mufns to
10%, whereas the molecular weights of the b-glucans were
similar for both fresh and frozen products. Blood glucose
response was determined in healthy subjects over 2 h
and the peak blood glucose rise (PBGR) was signicantly reduced after consuming fresh mufns compared to
those submitted to four freezethaw cycles (1.84 vs.
2.31 mmol/L), i.e. fresh mufns reduced the area under
the glucose response curve twice as much as mufns
submitted to the freezethaw process. Furthermore, in
mufns containing b-glucans of similar molecular weight,
there was a signicant inverse linear relationship between
log(concentration) of solubilised b-glucan and PBGR.
Similarly, Ostman et al. (2006) reported a useful tool for
prediction of the glycemic responses to bread products
enriched with b-glucans. These researchers found a high
correlation (r 0.98) between the GI of normal human
subjects after consumption of b-glucan-enriched breads
and the uidity index (FI) that was estimated in the in vitro
enzymatic digests of bread products using a Bostwick
consistometer. This relationship was described by the
following equation:
DG 7:93 0:68 log10 c 1:01 log10 M w r2 0:88.
4.2.3. Fate of b-glucan in the gastro-intestinal tract

Animal studies indicate a decrease of b-glucan molecular
weight during digestion. Thus, Wood and co-workers
demonstrated a signicant depolymerisation of oat
b-glucan during digestion in the small intestine of rats
and chicks (Wood, 1994; Wood et al., 1991a); the maximum b-glucan concentration was found in the ileal section,
where most of the starch has been absorbed. The molecular
weight of oat b-glucan extracted from the intestinal content
of rats was lower relative to that in the intact diet; a 10-fold
lower molecular weight of b-glucan was present in the
small intestine than in the stomach, whereas there were
further decreases in the caecum and faeces (Wood et al.,
1991c). Dongowski et al. (2002) found that b-glucan was
highly fermentable in the rat caecum and was not found in
faeces. The molecular weight of b-glucan from the stomach
and intestines of hamsters was about 100,000, lower than
that of bre subjected to processing treatments (Yokoyama
et al., 1998, 2002). Even in pigs, which are usually considered good models for humans, depolymerisation of
b-glucans to a molecular weight of about 100,000 occurred
during passage through the stomach and the upper jejunum
(Johansen et al., 1993). This depolymerisation was attributed to enzymic activity of microbes present in the gut,
but their action does not lead to a loss in b-glucan from the
digesta before it reaches the lower small intestine; trypsin
had no effect on the viscosity of b-glucan solutions. On the
other hand, Robertson et al. (1997) showed that endogenous proteases from the small intestine of ileostomate
(3)
However, real foods may differ greatly from the drink
model described above. Since the viscosity developed in the
gastro-intestinal tract seems to be the most critical variable
for the physiological effects of b-glucans, it is important to
also take into account the modication of the bre matrix
during its transit in the small intestine when assessing the
dietary response to a bre source. There is also some
evidence of changes in the solubility and molecular weight
of b-glucan during digestion. Beer et al. (1997b) extracted
b-glucan from oat bran and from oat bran mufns using an
in vitro system that included a sequential treatment with
amylase and pepsin to crudely mimic the human digestion
process. The original brans used, prior to mixing with other
ingredients and cooking, showed about 2530% b-glucan
extractability and a molecular weight of 12 106, while
cooking of the oat bran mufns resulted in reduction of the
molecular weight to 600950 103, but increased the
percentage of b-glucan solubilised by about threefold (to
5585%), depending on the recipe followed. During frozen
storage, the extractable b-glucan was reduced to 2740% in
all mufns, but no change in the peak molecular weight
of b-glucans was detected; the reduced solubility of the
b-glucan possibly reects changes in the molecular organisation (chain aggregation) during frozen storage. In a
recent study, Brummer et al. (2006) successfully used this in
vitro assay to predict the effectiveness of a solid food in
GI 50:8 0:441 FI:
(4)
ARTICLE IN PRESS
114
humans were capable of enhancing the extractability of the

barley b-glucan measured in vitro to a level similar to that
found in ileal efuent from the patients fed an acute barley
based-diet. The extractability measured in vivo was signicantly higher than that measured in the original barley
but without protease treatment. Increased extractability
can be accompanied by a reduction in the molecular weight
of the b-glucan in the upper gut, 10% of the total b-glucan
recovered had a molecular weight of o12,000. Similarly,
Sundberg et al. (1996) observed some depolymerisation of
b-glucans during the transit through the upper gastrointestinal tract of ileostomates consuming breads of
different cereals. Thus, unextractable b-glucans have the
potential to behave as a source of soluble bre because of
the modication during transit in the upper gut (Robertson
et al., 1997). It is also notable that the decrease in b-glucan
molecular weight was signicant only in breads containing
high molecular weight b-glucans, e.g. the 1.3 106 peak
molecular weight (Mwp) of b-glucan included in a oat
bread was found in excreta of ileostomic bags to be
0.7 106. In contrast, this reduction was limited for a
barley b-glucan of relative low molecular weight in the
consumed bread, i.e. the Mwp from 285 103 was reduced
to 262 103 after passage through the small intestine
(Sundberg et al., 1996).
4.2.4. Future perspectives
The ability of soluble bre to increase intestinal transit
time has been also attributed to the gel-formation properties of these materials (Anderson and Chen, 1986);
however, soluble bre such as oat bre may increase faecal
bulk and alleviate constipation (Malkki and Virtanen,
2001). The gel network that is formed could also act as a
molecular sieve in the small intestine; large molecules could
pass rapidly through the hydrated network, but smaller
molecules would be trapped in the various pores for
variable lengths of time slowing down the digestion
(retarded diffusion and contact between gastro-intestinal
enzymes and their substrates) and absorption of various
nutrients. However, the possible relationships between the
gelling properties of cereal b-glucans and their physiological functions have not yet been examined. To our knowledge, there are no reports in the literature addressing the
possible relationships between the structural features of bglucans and their physiological effects. The latter can be
modulated by interchain aggregation phenomena which
are controlled by the ratio of cellotriosyl/cellotetraosyl
units, the sequence of cellotriosyl residues along the chain,
and the presence and distribution of long cellulose-like
fragments. Further clinical studies must be properly
designed with well-characterised materials to provide
evidence for such structurefunction relationships.
5. Conclusions
The physical and physiological properties of b-glucans
are of commercial and nutritional importance. Recent
studies show that the molecular and structural features of

cereal b-glucans are important determinants of their
physical properties, such as water solubility, dispersibility,
viscosity, and gelation properties, and thereby affect the
functionality of these polysaccharides from a technological
viewpoint when added to food systems. The physiological
functions of cereal b-glucans also seem to be associated
with viscosity enhancement properties, and this may be
affected by the amount and molecular weight of solubilised
b-glucan in the gastro-intestinal tract. However, the
possible relationships between the gelling properties of
b-glucans and their physiological effects need to be further
studied.
Acknowledgements
This work has been carried out with the nancial support
from the Greek Ministry of Development, General
Secretariat for Research & Technology, International
Cooperation in Industrial Research and Development
Program DSEBPRO-2005, # 05 DSEBPRO-100, 20062008.
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Journal of Cereal Science 46 (2007) 101118

Molecular aspects of cereal b-glucan functionality: Physical properties,

Abbreviations: DP, degree of polymerisation; DP 3, 3-O-b-cellobiosyl-D-glucose; DP 4, 3-O-b-cellotriosyl-D-glucose; DPX5, cellodextrin-like

3.2.1. Dynamic rheometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

ward et al., 1983a, 1988). Cereal b-glucans exhibit

Long cellulosic unit (n3)

sieving/centrifugation and precipitation of the polysaccharide components with alcohol solution.

Wood et al. (1991ac)

Cui et al. (2000) and Li

Dais and Perlin, (1982)

attributed to genotypic and environmental factors (Jiang

varieties appear to have a higher DP3:DP4 ratio compared

aleurone tissues are higher than those from starchy

Burkus and Temelli 2003; Colleoni-Sirghie et al., 2003b;

where Z is viscosity, Z0 is viscosity at the Newtonian zone, g_

in the concentration of dissolved polysaccharide. This rule

et al., 2006; Papageorgiou et al., 2005; Skendi et al., 2003;

decreases and the IE values increase with decreasing

Furthermore, the dependence of the storage modulus

Fig. 5. Cereal b-glucan molecular weight and structure effects on tan d

Endothermic Heat Flow

2003, 2004), as well as the apparent melting enthalpy values

curing 3% (w/v) b-glucan dispersions at 5 1C for 135 days,

A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118

seemed to be a coarsening of the gel network structure as

characteristics and under certain conditions is a desirable

(Skendi et al., 2003). The mechanical properties of cast

inclusion of 5% of a b-glucan-rich fraction in bread

A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118

the cholesterol-lowering effect is larger in subjects with

It should also be noted that the doseresponse for a solid

effects on the concentrations of blood lipids, insulin or

attenuating the glycemic response; this was achieved in

DG 7:93 0:68 log10 c 1:01 log10 M w r2 0:88.

4.2.3. Fate of b-glucan in the gastro-intestinal tract

GI 50:8 0:441 FI:

A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118

humans were capable of enhancing the extractability of the

studies show that the molecular and structural features of

A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118

water-extractable b-glucans from different Greek barley cultivars.

with multiple-angle laser light scattering and other detectors. Cereal

endosperm. III. Distribution of cellotriosyl and cellotetraosyl residues.

A. Lazaridou, C.G. Biliaderis / Journal of Cereal Science 46 (2007) 101118

Wikstrom, K., Lindahl, L., Andersson, R., Westerlund, E., 1994.

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