J Nutr Sci Vitaminol, 64, 8–17, 2018

Review
b-Glucan in Foods and Its Physiological Functions

Ayaka Nakashima, Koji Yamada, Osamu Iwata, Ryota Sugimoto, Kohei Atsuji,
Taro Ogawa, Naoko Ishibashi-Ohgo and Kengo Suzuki*
euglena Co., Ltd., Tokyo 108–0014, Japan
(Received August 14, 2017)

Summary b-Glucans are a class of polysaccharides consisting of d-glucose units that are
polymerized primarily via the b-1,3 glycosidic bonds, in addition to the b-1,4 and/or b-1,6
bonds. They are present in various food products such as cereals, mushrooms, and seaweeds
and are known for their numerous effects on the human body, depending on their struc-
tures, which are diverse. The major physicochemical properties of b-glucans include their
antioxidant property, which is responsible for the scavenging of reactive oxygen species, and
their role as dietary fiber for preventing the absorption of cholesterol, for promoting eges-
tion, and for producing short-chain fatty acids in the intestine. Dietary b-glucans also exert
immunostimulatory and antitumor effects by activation of cells of the mucosal immune
system via b-glucan receptors, such as dectin-1. In this review, we elaborate upon the diver-
sity of the structures and functions of b-glucans present in food, along with discussing their
proposed mechanisms of action. In addition to the traditional b-glucan–containing foods,
recent progress in the commercial mass cultivation and supply of an algal species, Euglena
gracilis, as a food material is briefly described. Mass production has enabled consumption of
paramylon, a Euglena-specific novel b-glucan source. The biological effects of paramylon are
discussed and compared with those of other b-glucans.
Key Words b-glucan, food material, paramylon, food functionality

b-Glucans as Food Constituents
b-Glucan is a nonstarch polysaccharide that con-
sists of d-glucose units linked via b glycosidic bonds.
Typically, b-glucans do not include cellulose, which is
formed only by b-1,4 glycosidic bonds. So-called b-glu-
can is a polysaccharide formed mainly via b-1,3 glyco-
sidic bonds, with varying numbers of b-1,4 and b-1,6
glycosidic bonds. The cell walls and aleurone layers of
various organisms are made of these b-glucans. There-
fore, human beings unintentionally consume b-glucans
through their regular diet. The primary sources of
dietary b-glucan for humans are cereal grains (espe-
cially oats and barley), mushrooms, seaweeds, and yeast.
The functions of b-glucan have been recognized for
more than half a century. In pioneering studies, zymo-
san, which was prepared from yeast cell walls, has been
found to contain a high percentage of b-glucan (1) and
has been reported to cause nonspecific activation of the
immune system (2). On the basis of the accumulated
knowledge related to b-glucan functions, lentinan and
sonifilan prepared from mushrooms have been used as
b-glucan-based medicines for their immunostimulatory
effects (3, 4). Although the b-glucan–based medicines
are administered via injections, oral administration
of some b-glucans has been reported and has similar
immunostimulatory effects. In addition, the dietary
b-glucans present in cereals (such as oats and barley)
* To whom correspondence should be addressed.
E-mail: suzuki@euglena.jp
have been demonstrated to reduce the level of serum
cholesterol and to act as major dietary fiber prevent-
ing obesity and metabolic disorders (5, 6). The reported
functions of dietary b-glucan have encouraged the con-
sumption of b-glucan–rich foods and the development
of several functional foods that contain b-glucans.
Types of b-Glucans
b-Glucans extracted from different organisms pos-
sess characteristic sizes; numbers of b-1,3, b-1,6, and
b-1,4 glycosidic bonds; and b-1,6 branching patterns
(Fig. 1, Table 1). Nonetheless, the size of a b-glucan
also depends on the method of extraction and purifica-
tion. The b-glucans present in cereals (such as oats and
barley) include a mixture of b-1,3 and b-1,4 glycosidic
bonds and show no b-1,6 branching (6, 7). On the
other hand, the b-glucans present in other organisms
contain fewer b-1,4 glycosidic bonds. The b-glucans
present in edible mushrooms [such as Lentinus edodes
(shiitake mushroom), Grifola frondosa (maitake mush-
room), and Schizophyllum commune (suehirotake mush-
room)] consist of a linear b-1,3-glucan backbone with
a single b-1,6-linked glucose unit present on two resi-
dues up to every third residue of the backbone (8–11).
In contrast, the b-glucan in zymosan, which is extracted
from the cell wall of Saccharomyces cerevisiae, is a large
molecule consisting of a linear b-1,3-glucan backbone
with a 30-residue linear side chain (held together by
b-1,3 glycosidic bonds) attached to the main chain via
a b-1,6 glycosidic bond (12). The b-glucans present in

8
 b -Glucan-Containing Foods

Fig. 1. Representative structures of b-glucans extracted from cereals, mushrooms, yeast, seaweeds, bacteria, and Euglena. (A) A cereal b-glucan consisting of consecutive b-1,3 and b-1,4
glycosidic bonds without b-1,6 branching. (B) A mushroom b-glucan consisting of a linear b-1,3-glucan chain with single b-1,6-linked glucose units attached to the backbone. (C) A
yeast b-glucan extracted as a large molecule showing a highly branched structure. It contains only b-1,6 glycosidic bonds in the b-1,3-glucan chain. (D) A seaweed b-glucan composed
of b-1,3-glucan chains with b-1,6 branching. Some seaweed b-glucans are characterized by the presence of mannitol at the end of the b-1,3-chain. (E) Bacterial and Euglena’s b-glucans
9

consisting of linear b-1,3-glucan chains without branching.

10 Nakashima A et al.

Table 1. A summary of physicochemical properties of representative b-glucans present in food products.

Name of the Example

Category General name Species Glycosidic bond(s) Solubility Reference
glucan of size

1.6 MDa,
Higher plant Oats Avena sativa — Soluble 7)
b-1,3 and b-1,4 glyco- 1.1 MDa
sidic bonds without
Hordeum branching
Higher plant Barley — 49 MDa Soluble 6)
vulgare

Maitake
Mushroom Grifola frondosa Grifolan 30 kDa1 Soluble 9)
mushroom
b-1,3 bonds present in
Shiitake the main chain with
Mushroom Lentinula edodes Lentinan 500 kDa Soluble 8, 11)
mushroom b-1,6-linked single
glucose units
Schizophyllan
Mushroom Schizophyllan SPG 450 kDa Soluble 10, 11)
commune

Laminaria
Seaweed Kelp Laminarin 4 kDa2 Soluble 15)
digitata

Saccharina
Seaweed Kelp Laminarin b-1,3 bonds present in 2.9–3.3 kDa Soluble 14)
longicruris
the main chain with
b-1,6-linked side
Durvillaea
Seaweed Cochayuyo Laminarin chains 6.9 kDa Soluble 13)
antarctica

Saccharomyces
Yeast Baker’s yeast Zymosan 240 kDa Insoluble 12)
cerevisiae

b-1,3-glucan chain
Microalgae Euglena Euglena gracilis Paramylon 500 kDa Insoluble 17, 18, 21)
without branching
1
Size of b-glucan in the MT-2 fraction.
2
Estimated by the polymerization degree of glucose.

Fig. 2. Microscopic images of (A) Euglena gracilis cells and (B) paramylon granules. Scale bars represent the size indicated
in the figures.

seaweeds such as Laminaria digitate, Saccharina longi-
cruris, and Durvillaea antarctica are reported to consist
of a b-1,3-glucan backbone too, with a small percent-
age of b-1,6 glycosidic bonds in the backbone, and
b-1,6-glucan side chain branching (13–15). The b-glu-
cans extracted from seaweeds are relatively smaller in
size compared to the b-glucans of other organisms and
include a monosaccharide other than glucose (like man-
nitol) at the end of the chain.
Among various b-glucans, including those men-
tioned above, the b-glucan with the simplest structure
is the linear and unbranched b-1,3-glucan curdlan,
which is extracted from the bacterium Agrobacterium
biobar (16). Although the organisms containing cur-
dlan are not edible, purified curdlan serves as a thick-
ening agent for food products. Nonetheless, the recent
commercialization of Euglena gracilis production has
provided an alternate source of linear and unbranched
 b -Glucan-Containing Foods 11

Fig. 3. Schematic representation of the mechanisms of recognition of pathogens and b-glucans in the gut. Both b-glucan
particles and pathogens have b-glucan residues on their surface. Both presumably pass through the epithelial lumen by
active transport mediated by M cells in Peyer’s patches; however, some soluble b-glucans may also be absorbed like other
nutrients. b-Glucans act on receptors expressed on leukocytes, such as macrophages and dendritic cells.

b-1,3-glucan. Euglena, which is a genus comprising
photosynthetic flagellates living in fresh and brackish
water, is known to accumulate b-1,3-glucan in its body
as a carbohydrate source (Fig. 2). Paramylon, i.e., crys-
tallized b-glucan from Euglena, has characteristic fea-
tures such as the presence of unbranched b-1,3-glucan
(17, 18), highly crystalline structure with a triple-helix
conformation (19, 20), and unique saccharide compo-
sition (exclusively consisting of glucose) (21). Owing to
its crystalline structure, paramylon typically exists in
the form of insoluble granules that are 2–3 mm in size.
The granules can be easily obtained by centrifugation
from cells disrupted by sonication and subsequently can
be purified by denaturing the proteins with a suitable
detergent (22). These features differentiate paramylon
from other b-glucans in that systematic analysis of its
functions is possible.
The types and strength of physiological functions of
b-glucans depend on their structure, such as composi-
tion of the glucan backbone, types and frequency of
side chains, size of the molecule, solubility, and three-
dimensional conformation. b-Glucans show viscosity
in an aqueous solution and thus function as dietary
fiber by adsorbing unnecessary materials in the gut and
helping to egest them (23). They also scavenge reactive
oxygen species (ROS) in the gut, thereby exerting an
antioxidant activity (24). These activities are common
for all b-glucans although they vary depending on the
physicochemical properties of each b-glucan species. In
addition, the structure of the b-1,3-glucan backbone
and b-1,6 branching at appropriate sites enhance the
immunostimulatory effect of b-glucans by promoting
their interaction with specific receptors (25, 26).
Dectin-1, a C-type lectinlike pattern recognition
receptor present on the surface of leukocytes, is known
as the primary receptor for b-glucans (27). The triple-
helix structure formed by the b-1,3-glucan backbone
is specifically recognized by dectin-1. Additionally, the
branches located on the surface of the helix further mod-
ulate this recognition (28, 29). An assay of the binding
of recombinant dectin-1 to various b-glucans indicates
a wide range of binding affinity (IC50, half maximal
inhibitory concentration, as 2.6 mm to 2.2 pm), depend-
ing on the size and branching patterns of b-glucans
(30). The interaction of dectin-1 with cereal b-glucan,
which includes both b-1,3 and b-1,4 glycosidic bonds,
has been confirmed; however, the binding affinity is
weaker than that for b-1,3-glucan (31). Another impor-
tant b-glucan receptor is complement receptor 3 (CR3),
which is a heterodimeric transmembrane glycoprotein
found on leukocytes (32). CR3 is known for respond-
ing to small soluble b-glucan molecules although it is
also activated by insoluble b-glucan to some extent (33,
34). In addition, b-glucans are reported to interact with
such receptors as langerin (35), lactosylceramide (36),
and L-ficolin (37), thereby regulating their respective
downstream signaling pathways (25). The receptor-
mediated effects of b-glucans are well characterized
for the b-glucan extracts of mushrooms and yeast that
serve as medicines. The b-glucan present in mushrooms
has considerable receptor-mediated effects. b-Glucans
extracted from other organisms exert the same effects
probably to a lesser extent (5, 6, 38). The differences in
these receptor-mediated functions of b-glucans seem to
depend on their intestinal uptake efficiency and affinity
for their receptor(s).
Functions of Dietary b-Glucans
For understanding the correlation between b-glucan
structure and function, the biological effects of dietary
12 Nakashima A et al.
b-glucans have been evaluated after their purification.
The differences in the b-glucan preparation proce-
dures under specific experimental conditions result in
observation of a limited range of effects. Thus, the real
potential of foods containing b-glucans is not necessar-
ily restricted to the reported effects. In addition, owing
to the difficulties with preparing highly pure and stan-
dardized b-glucans, careful evaluation excluding the
effects of contaminants is required for elucidation of the
true effects of dietary b-glucans. Various effects on the
human body after consumption of dietary b-glucans
have been reported, including a reduction in the cho-
lesterol level, prevention of diabetes, regulation of the
gut microflora, antioxidant activities, antitumor effects,
and immunostimulation. Some important effects are
discussed below.
a) Immunostimulation
The immune system is a framework for defending the
body against bacterial and viral infections (and other
invaders). The age-related reduction in immunological
function is referred to as immunosenescence, which
results in the increased susceptibility to infections in
the elderly. One of the primary causes of immunosenes-
cence is the age-related decline of T-cell function and
involution of the thymus (39, 40). Ingestion of b-glucan
exerts a stimulatory effect on immune homeostasis via
the b-glucan receptor present in the mucosal immune
system and contributes to the prevention of age-related
diseases (41). This effect has been reported for almost all
b-glucan–containing foods although the effect size may
vary (5, 33, 38, 42).
The immunostimulatory effect of b-glucans is medi-
ated by innate immunity intended for the defense
against pathogens (43). Upon ingestion, b-glucans
affect the mucosal immune system in the gastrointes-
tinal tract (Fig. 3). The uptake of microorganisms from
the intestinal lumen is undertaken by the M cells of Pey-
er’s patches in the small intestine. Subsequently, these
cells present the antigen (or the microorganism itself)
at their basal surfaces to immune cells such as macro-
phages and dendritic cells (44). To identify the patho-
gens, the receptors specific for pathogens’ cell wall are
enriched on the surface of immune cells. Several pattern
recognition receptors target b-glucans that are present
in the cell wall of microorganisms. In addition to act-
ing as a stimulatory signal on cell surface receptors, the
microorganisms that pass through the intestinal lumen
are phagocytosed by macrophages and have additional
immunostimulatory effects. Microparticles that include
insoluble b-glucan and range in size from 1 to 2 mm
show strong immunostimulatory effects (45), probably
because of being recognized as pathogens. Although it is
not clear how b-glucans pass through the epithelial cell
lining, some epithelial cells such as M cells may actively
absorb b-glucans (like other antigens). The difference in
the immunostimulatory effects of insoluble and soluble
b-glucans suggests that they act via different molecular
mechanisms (33). Moreover, purified b-glucans, such
as lentinan, which are generally administered via injec-
tion, have a considerably smaller immunostimulatory
effect after oral administration (46), thereby indicating
that b-glucans also act differently depending on their
route of administration.
Dectin-1, which is expressed on the surface of mac-
rophages and dendritic cells, is reported to serve as the
primary receptor for b-glucans, which consequently
have immunostimulatory effects (27). Depending on the
recognition of b-glucan by dectin-1, those cells phago-
cytize particles displaying b-glucan on their surface as
well as the pathogens that include b-glucan in their cell
wall (47). In parallel with the phagocytosis, dectin-1
induces secretion of proinflammatory cytokines (48,
49) via the activation of tyrosine kinase Syk and a tran-
scription factor, nuclear factor kB, which is responsible
for multiple immune reactions (33). Insoluble b-glucan
preferentially binds to dectin-1, whereas water-soluble
b-glucans bind to another receptor, CR3, and thus trig-
ger an immune reaction (33). Meanwhile, micronized
and water-soluble b-glucans are probably also absorbed
by the small intestine following ingestion, reach as far
as the circulatory system, and increase the expression
levels of dectin-1 and a Toll-like receptor 2 (TLR2),
which together with dectin-1 recognizes b-glucan in
gut-associated lymphoid tissue (50). Thus, it seems that
b-glucans are absorbed in the body and act in different
ways to elicit an immune reaction depending on their
structure.
b) Immune regulation of the T helper 1 (Th1)/Th2 balance
The importance of balanced immune homeostasis
has been recognized in recent years because excessive
immune responses can lead to autoimmune diseases or
allergies (51). One example of immune homeostasis is
the balance between Th1 cells of cell-mediated immu-
nity and Th2 cells of humoral immunity (52). The Th1-
and Th2-mediated immune reactions suppress each
other for immunological homeostasis. Atopic dermatitis
represents an example of allergic reactions caused by an
imbalance between Th1 and Th2 cells. An alteration of
the immunological equilibrium in the Th2 direction is
responsible for the symptoms of atopic dermatitis and
other allergies (53). Meanwhile, excessive activation of
the immune system mediated by Th1 cells together with
that of Th17 cells causes autoimmune diseases, such
as rheumatoid arthritis (54). Ingestion of b-glucans
primarily activates Th1 and Th17 cells (55), thereby
resulting in the suppression of activity of Th2 cells. This
phenomenon shifts the Th1/Th2 response equilibrium
in the Th1 direction and alleviates the symptoms of
allergic diseases (56). This action is basically triggered
by the immunostimulatory effect of b-glucans, sug-
gesting that all b-glucan–containing foods possessing
immunostimulatory effects also have this activity.
c) Cholesterol-lowering property
The cholesterol-lowering property of the dietary
b-glucan of cereals, such as oats and barley, has been
known for several decades and has been verified clini-
cally by prescription of dietary b-glucan to patients with
high levels of serum cholesterol (5, 6). The lowering of
cholesterol levels does not affect high-density lipopro-
tein (HDL) cholesterol; however, it affects low-density
 b -Glucan-Containing Foods
lipoprotein (LDL) cholesterol, which is an important
marker of the risk of cardiovascular diseases (57, 58).
This means that consumption of dietary b-glucans pres-
ent in cereals lowers the risk of cardiovascular diseases.
For example, consumption of bread formulated with
oat b-glucan significantly lowers LDL-cholesterol lev-
els (59). Because the cholesterol-lowering property has
only been reported for the b-glucans derived from cere-
als, the effects of other b-glucan-containing foods on
cholesterol levels are uncertain at present.
The reduction in cholesterol levels by b-glucan is
primarily due to its ability to act as dietary fiber and to
entrap the bile acid micelles containing fats. This activ-
ity disrupts the interaction of the micelles with lumi-
nal membrane transporters present on the intestinal
epithelium, thereby increasing the fecal output of fat,
bile acids, and cholesterol (23). Next, to compensate
for the decrease in the bile acid level, the expression of
7a-hydroxylase (an enzyme involved in the synthesis
of bile acids) is upregulated (60, 61). Accordingly, the
hepatic cholesterol level also decreases and is restored
by LDL-cholesterol from serum. In addition, the delayed
absorption effect due to the high viscosity of b-glucans
from cereals may lower postprandial glucose levels and
prevent diabetes (62).
d) Fermentation effects of b-glucan in the colon
Soluble b-glucans, especially those present in cere-
als, are fermented by the microorganisms residing in
the colon and are converted to short-chain fatty acids
(SCFAs) such as acetic acid, propionic acid, and butyric
acid (63). The SCFAs produced in the colon have various
effects, such as immunomodulation (64), mediation of
apoptosis of colon cancer cells (65), and prevention of
obesity (66). As an example, the prevention of obesity is
accomplished by regulation of energy metabolism and
prevention of excess lipid accumulation in adipose tis-
sue via SCFA receptor GPR43 (66). Besides, SCFAs sup-
press the growth of harmful bacteria such as Clostrid-
ium spp. and pathogenic coliforms and thereby maintain
a healthy gut microflora (67).
e) Antioxidant activity
Cereals are known to exert antioxidant activities by
scavenging ROS. In particular, the ROS-scavenging abil-
ity of oats and barley mainly derives from their abun-
dant b-glucans, which are known to scavenge ROS as
effectively as other polysaccharides do (5, 6). The b-glu-
can extracted from barley shows a significantly higher
antioxidant activity as compared to b-glucan from oats
and black yeast, indicating that the structure and com-
position of b-glucans also influence their antioxidant
activity (24). Meanwhile, the ROS-scavenging activity
of b-glucan–containing foods other than cereals may
be low because they include lesser amounts of b-glu-
can, and its abundance is important for this action. The
antioxidant effects of ingested cereal b-glucan possibly
prevent the damage to the gut that is caused by ROS
generated there. In addition to the reports on the physi-
cal ability of b-glucan to scavenge ROS, several stud-
ies on rats as a model organism have revealed that the
antioxidant activity of orally administered b-glucan is
13
also directed against oxidative stress induced even in the
internal organs such as kidneys and the liver (68, 69).
These reports imply that b-glucan also exerts the anti-
oxidant effect by either of the following mechanisms:
b-glucan incorporated into the circulatory system
scavenges ROS there, or b-glucan stimulates the innate
antioxidant system via specific receptors in the mucosal
immune system and the subsequent cytokine release.
f) Antitumor activity
The medicinally important b-glucans—lentinan, gri-
folan, and schizophyllan glucan (SPG), which are puri-
fied from shiitake mushrooms, maitake mushrooms,
and Schizophyllum, respectively—are used as antitu-
mor drugs (3, 4). The antitumor effects are attained
via intravenous administration (70). Nevertheless, oral
administration of the recently developed superfine dis-
persed lentinan has shown significant effects on hepa-
tocellular and pancreatic cancers (71, 72). This finding
suggests that eating mushroom fruiting bodies itself
may have potential antitumor effects. The antitumor
activity of b-glucans via the b-glucan receptors is pri-
marily mediated by an increase in the tumor immunity,
which enhances the killing of iC3b-opsonized tumor
cells (33). Consistent with this observation, the antitu-
mor activities of other b-glucans, which are known for
their immunostimulatory effects, have been reported
as well (5, 33, 73). In addition to their effect on tumor
immunity, the antitumor activity against colon cancer
may be attributed to other factors such as induction of
egestion of a carcinogen in the gut, regulation of the
gut microflora, and scavenging of ROS, as mentioned in
subsections (c), (d), and (e), respectively.
Paramylon as a Functional Ingredient of Food
Due to the establishment of commercial mass cultiva-
tion of E. gracilis in 2005, a reliable and economically
feasible supply of purified paramylon and the algal pow-
der containing large amounts of paramylon is now pos-
sible (74). Because paramylon has a crystalline struc-
ture composed of b-1,3-glucan chains, it presumably
has effects on the human body that are similar to those
of other b-glucans. Thus, it can serve as a substitute or
supplement for b-glucan–based applications. To evalu-
ate the effects of ingestion of paramylon compared to
that of other b-glucans, a series of experiments have
been conducted.
To determine whether paramylon has an antioxidant
activity, its effects on oxidative liver damage induced by
carbon tetrachloride have been examined in rats (75).
It was observed that the carbon tetrachloride–induced
damage is suppressed after 3 d of oral preadministra-
tion of paramylon, indicating that paramylon has an
antioxidant activity. The effects of oral administration
of paramylon on oxidative liver damage suggest that
it also possibly prevents oxidation-induced damage to
other organs by a mechanism similar to that of other
b-glucans (68, 69).
The dectin-1–mediated immunostimulatory effect of
paramylon in animal disease models has been reported
too. For example, oral administration of paramylon to
14 Nakashima A et al.
the rainbow trout has been found to be effective against
the enteric redmouth infection, a bacterial disease of
salmonid fishes (76). In addition, paramylon enhances
antibody responses in mice against sheep red blood
cells by inducing the production of cytokines IL-1 and
IL-6, which were primarily released by macrophages
(77). A similar beneficial effect of paramylon has been
demonstrated in a mouse model of atopic dermatitis
(78). In contrast to the immunomodulatory effect of
the shiitake mushroom-derived b-glucan lentinan via
shifting the Th1/Th2 response equilibrium toward the
Th1 response (79), paramylon suppresses both Th1 and
Th2 cell responses in a mouse model of atopic derma-
titis (78). This finding suggests that paramylon has an
additional function as compared to the other b-glucans.
The structural differences between paramylon and the
other b-glucans may be responsible for varying levels
of receptor activation and specificity. Given that oral
administration of paramylon also exerts an immuno­
stimulatory effect in healthy human subjects (80), it is
capable of adequately regulating immune responses.
The antitumor activity of paramylon has been evalu-
ated by testing the suppression of colon cancer aberrant
crypt foci (ACF; precancerous lesions that form before
colon polyps and are detected at the earliest stage of
colon cancer) in mice. After paramylon treatment, ACF
formation mediated by administration of the chemi-
cal carcinogen 1,2-dimethylhydrazine is suppressed by
59% (81), indicating that paramylon exerts an antitu-
mor activity. Just like dietary b-glucans present in cere-
als, paramylon has an anti-ACF preventive effect that is
potentially mediated by a combination of tumor immu-
nity enhancement, antioxidation, and egestion activities
(5, 6). Due to the unique crystalline structure of par-
amylon granules purified from E. gracilis, they cannot
be hydrolyzed by any b-1,3-glucanases (82). This char-
acteristic feature differentiates paramylon from other
b-glucans. In the absence of intestinal fermentation of
paramylon, it is expected to have long-lasting effects.
Furthermore, the hydrolysis resistance of paramylon
may be partly responsible for its effects on the intes-
tine as a dietary fiber. This phenomenon increases the
fecal weight and shortens the intestinal residence time
for contents of the digestive tract (83). Additionally, it
adsorbs harmful carcinogens and promotes their excre-
tion (81), thereby possibly preventing colon cancer.
The effects of paramylon are similar to those of other
dietary b-glucans. This observation suggests that par-
amylon also functions by being transported across the
epithelial cells, is recognized by macrophages via recep-
tor dectin-1, and induces a Th1 cell response. The size
of a paramylon molecule is typically 2–3 mm (Fig. 2B),
which is similar to the size of pathogenic bacteria. This
observation suggests that paramylon is potentially
transported across the epithelial cells via the same
mechanism as that for pathogenic bacteria. The crys-
talline structure of paramylon consists of triple helices
(19, 20), which are recognized by b-glucan receptor
dectin-1, implying that paramylon presumably activates
dectin-1. In agreement with the above data, paramylon
directly binds to recombinant dectin-1 (84) and upregu-
lates proinflammatory factors such as NO, TNF-a, IL-6,
and COX-2 (80). Although studies have suggested that
the number and size of b-1,6 branches on the b-1,3
backbone are important for the function of b-glucan
(25, 26), paramylon, despite limited branching, shows
functional diversity. This phenomenon indicates that
paramylon derived from Euglena may serve as a useful
food ingredient to provide benefits similar to those of
other dietary b-glucans.
Conclusion
The physiological functions of dietary b-glucan can
be subdivided into two groups depending on whether
specific receptors are involved or not. Functions that do
not depend on specific receptors include the ROS-scav-
enging activity (antioxidant activity) resulting from the
physicochemical properties of b-glucan and the ability
to lower the level of serum cholesterol and to improve
the intestinal bacterial flora as dietary fiber. Such func-
tions are reported mainly for cereal b-glucans. Mean-
while, specific receptors such as dectin-1 and CR-3 are
involved in immunoregulatory functions and the anti-
tumor activity, which are well known for mushroom
b-glucans. In addition, relatively strong immunomod-
ulatory and antitumor effects have been reported for
Euglena paramylon, which was recently validated as a
functional food ingredient.
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