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Beta-glucan

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DOI: 10.1016/B978-0-323-89779-2.00013-2

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Chapter 20

Beta-glucan
Hanuman Bobade, Antima Gupta and Savita Sharma
Department of Food Science and Technology, Punjab Agricultural University, Ludhiana, Punjab, India

20.1 Introduction
Beta-glucan is a predominant non-starch polysaccharide composed of linear chains of b-D-glucose linked by b-(1 / 3),
(1 / 4), and/or (1 / 6)-D-glycosidic linkage consisting of over 25,000 D-glucose units either in branched or unbranched
form (Ahmad & Kaleem, 2018; Kaur et al., 2019). Glucans may be classified into two types: (a) water-soluble glucans and
(b) water-insoluble glucans. Salecan and curdlan are the examples of water-soluble and water-insoluble glucans,
respectively. Molecular mass, composition, crude preparation, and chemical structure could affect the solubility of beta-
glucan (Xiu et al., 2011).
Beta-glucan is one of the important constituents of cell walls of most fungi and many plants. The major source of beta-
glucan involves cereals, microorganism, mushroom, lichens, and seaweeds. The beta-glucan content differs largely among
the various sources. Among the cereals, greater amount of beta-glucan is found in barley followed by oat while least
amount is found in wheat and rice (Mejía et al., 2020). Beta-glucan found in cereal brans like barley and oat is usually
produced as agricultural by-products. Extraction and isolation of beta-glucans from such sources imply their economic and
environmental benefits (Chen et al., 2011).
Beta-glucan is known to associate with its prebiotic effects leading to numerous health benefits such as maintaining
serum cholesterol, blood glucose levels, cardiovascular diseases, hypertension, cancer, and immune-enhancing properties
(Jayachandran et al., 2018; Kaur et al., 2019; Volman et al., 2008). Many of these physiological activities of beta-glucan
are attributed to its high viscosity (Mejía et al., 2020), which depends on molecular weight of the beta-glucan. It has been
recommended by US Food and Drug Administration, ingestion of 3 g per day of beta-glucan alongside low cholesterol diet
reduces the risk of cardiovascular diseases.
Beta-glucan content and its properties like solubility, viscosity, molecular weight are affected by different processing
conditions (Kivelä et al., 2009; Tosh et al., 2010). Solubility and molecular weight of beta-glucan are reduced during
cooking and freezing storage (Lovegrove et al., 2000). Many studies conducted on safety of beta-glucan suggested that
beta-glucan is safe to consume and is not associated with any adverse health effects (Babícek et al., 2007; Jonker et al.,
2010; Delaney, Carlson, Frazer, et al., 2003; Delaney, Carlson, Zheng, et al., 2003). Consumption of concentrated beta-
glucan sourced from barley had no toxic effects in rats and mice even on consumption of large amount of beta-glucan
(Delaney, Carlson, Frazer, et al., 2003; Delaney, Carlson, Zheng, et al., 2003). The beta-glucan toxicity was not
observed at fairly large concentration of 2000 mg per kg body weight (Delaney et al., 2004).

20.2 Sources/Derivatives
Since ages, cereals like oat, barley, and wheat are the recognized source of beta-glucan. Moreover, with expansion in
research, beta-glucans have also been extracted from microorganism (yeast, fungi, bacteria), mushroom (Pleurotus spp.
and Lentinula spp.), lichens (Cetraria islandica), and seaweed/algae (Laminaria spp. and Phytophthora spp.) (Ahmad
et al., 2018). Fig. 20.1 outlines different sources of beta-glucan. In cereals, beta-glucan is present in the aleurone layer
and endospermic portion while in microorganism it is present in the outer cell wall (Bernstein et al., 2013; Fesel &
Zuccaro, 2016).
The content of beta-glucan is higher in barley than in any other cereal ranging from 5.0 to 11.0% and it is present in
endospermic and aleuronic layers. The percentage of beta-glucan in oats ranges between 4.5% and 5.5% and it is mainly

Nutraceuticals and Health Care. https://doi.org/10.1016/B978-0-323-89779-2.00013-2

Copyright © 2022 Elsevier Inc. All rights reserved. 343
344 Nutraceuticals and Health Care

FIGURE 20.1 Different sources of beta-glucan.

present in the aleuronic and subaleuronic layers. The content of beta-glucan is relatively low in wheat ranging between
0.2% and 1.2%. Major portion of beta-glucan in wheat is present in wheat bran, thereby wheat milling by-products are
mainly utilized as the major source for beta-glucan extraction. In mushroom, it varied between 3.1% and 46.5%. In
mushroom, beta-glucan is mainly concentrated in mycelia and/or fruiting bodies. In yeast, it ranges between 5% and 7%
and found mainly in the outer cell wall of yeasts like Saccharomyces cerevisiae, Debaryomyces hansenii, Kloeckera
apiculata, Candida milleri, Schizosaccharomyces pombe, Zygosaccharomyces bailii, and Pichia membranifaciens.
Bacteria are also the potential source of beta-glucan; during fermentation a linear curdlan structure is produced by
Agrobacterium, Bacillus, and Pseudomonas spp.

20.3 Extraction and characterization techniques

Several processes and raw material are available for the extraction of beta-glucan but latest research is prioritizing on the
nonaggressive method of extraction and purification so that minimal changes will occur in glucan structure. A range of
extraction and purification methods are available for beta-glucan including solvent extraction, hot water extraction, alkali
extraction, enzymatic extraction, and ultrasound/microwave-assisted extraction. The methods are used singly or in com-
bination with each other for better recovery. Fig. 20.2 represents the general details of beta-glucan extraction.

20.3.1 Extraction of beta-glucan from cereals

There are two major techniques of separating beta-glucan from cereals named dry and wet separation techniques. In dry
separation technique, pearling and air classification methods are employed because of its simplicity. The recovery by dry
separation technique is usually less than 30% (Zhang et al., 2015) where wet processing results in 50%e70% recovery.
Whole cracked barley/oats/wheat, meal, or flour was hydrolyzed using acidified water or alkaline solution to solubilize
beta-glucan followed by centrifugation and ethanol precipitation (inhibit some Indigenous enzymes, remove dissociates
sugars, proteins, and some nonpolar compounds). After crude extraction, beta-glucan is purified using freezing-thawing
and alcohol method and removes starch, protein, fat, and many other unwanted species. The extractability of beta-
glucan depends upon particle size, pH, temperature, extraction time, solvent, and flour ratio. During extraction and pu-
rification, an additional step of inactivating b-glucanases should be employed. This enzyme is responsible for hydrolyzing
beta-glucan that results in altering the structure, molecular weight, and viscosity of the beta-glucan, as a result its
cholesterol lowering and glucose modulating activity reduces. Enzyme inactivation could be possible by autoclave, HCl,
 Beta-glucan Chapter | 20 345

FIGURE 20.2 General details of beta-glucan extraction.

trichloroacetic acid, ethanol precipitation, and heating. The basic disadvantage with wet separation technique is its high
cost of production. Addition to enzyme inactivation, starch removal is another problem of concern which can be removed
by water, acid, and alkali extraction (Song et al., 2008). It was observed by many researchers that beta-glucan is degraded
by acid, thereby alkali and hot water extraction is generally used to remove starch. In purification, dialysis and ultrafil-
tration technique is used, but former is little time-consuming process and is usually avoided at commercial level. Wang &
Zhao (2007) reported isoelectric precipitation method for purifying crude beta-glucan and observed purity was up to
91.38% with relatively low yield. The process in the form of flow sheet for extraction of beta-glucan from oat is indicated
in Fig. 20.3.

20.3.2 Extraction of beta-glucan from yeast

Yeast is one of the known sources of beta-glucan; it contains around 5%e7%. It is mainly present in the cell wall structure
of yeast and was first rendered into soluble form by alkaline extraction. Suphantharika et al. (2003) optimized the alkaline
extraction parameters including extraction time, 1 h; extraction temperature; sodium hydroxide (NaOH) concentration, 1N;
and yeast cell wall to NaOH ratio, 1:5 (w/v). Crude protein isolated was purified using different techniques such as DEAE-
cellulose, ConA chromatography, gel filtration chromatography, etc. The generalized procedure for isolating glucan from
yeast involves breakdown of cells by chemical, biochemical, mechanical, and autolysis method followed by separation of
cell wall using centrifugation and/or filtration. The process of this method is illustrated through flow sheet in Fig. 20.4.
Thereafter, beta-glucan is extracted by alkaline or any suitable extraction technique and later purified either by chro-
matographic technique or by precipitation and centrifugation method.
Generally, autolysis method is used for the lysis of cell. It is an irreversible process, caused by intracellular enzymes
chiefly b-(1 / 3)-glucanase and proteases, under stress conditions like pH, temperature, and yeast concentration which are
not suitable for the survival of cell. For better separation a thermal shock (60 C for 15 min) used to be given prior to
autolysis. Lysed cell wall is separated and beta-glucan is extracted by hot alkaline extraction followed by purification by
chromatographic method and/or precipitation method.

20.3.3 Extraction of beta-glucan from fungi

Beta-glucan extraction using hot water is the most common and convenient method of extraction. Several researchers
extracted beta-glucan from the fruiting bodies of fungi such as lentinan from Lentinus edodes (Bhanja et al., 2014),
schizophyllan from Schizophyllum commune (Kumari et al., 2008), Agaricus blazei (Yea-Woon et al., 2005), Ganoderma
346 Nutraceuticals and Health Care

FIGURE 20.3 Extraction of beta-glucan from oats.

FIGURE 20.4 Process for extraction of beta-glucan from yeast.

 Beta-glucan Chapter | 20 347

lucidum (Pai-Feng et al., 2012), etc. Isolation and extraction of beta-glucan from fruiting bodies are done by hot or boiling
water for around 3e4 h and later on crude beta-glucan is purified using column chromatography (Sephadex G-50 and/or
Sephadex A-25) and gel filtration chromatography (Sephadex G-15) techniques.
High-performance anion-exchange chromatography assisted with enzymatic treatment (lichenase) has been widely used
for the quantitative structural analysis of beta-glucan, in which exact amount of DP3 and DP4 could be estimated (Ryu
et al., 2009; Wood, 2011). FT-IR analyzes the beta-glucan structure very rapidly and sensitively. NMR is another widely
used technology to analyze the overall structure and linkage sequence in beta-glucan. 1H-NMR and Raman spectroscopy
are used to identify the compositional and structural features of beta-glucan which could further linked with its practical
ability and physiological impact (Mikkelsen et al., 2010). The absorption of beta-glucan in Raman spectroscopy is at 890/
cm where in case of 1H-NMR the resonance of b-(1 / 3) and b-(1 / 4) was observed at 4.75 and 4.55 ppm spectra
(Salomonsen et al., 2008). A range of methods like physical (low or high temperature), chemical processes (acid, alkali,
and salt), mechanical (homogenization, ultrasonication, etc.), and enzymatic treatment (glucanase, lichenase, etc.) are
employed to modify the beta-glucan in order to exploit the properties and activities of beta-glucan.

20.4 Chemistry
As depicted in Fig. 20.5, the beta-glucan is a non-starch polysaccharide consisting of linear chains of D-glucose having
(1 / 3)-b-D-glycosidic linkage with side chains attached to b-(1 / 6) in fungi and b-(1 / 4) in plants and bacteria
(Chan et al., 2009; Magnani & Castro-Gómez, 2008). In oats, unbranched linear structure of b-(1 / 3) and b-(1 / 4)-D-
glycosidic linkage is present in random fashion. It comprises 70% of b-(1 / 4) and 30% of b-(1 / 3)-glycosidic bonds in
its structure (Wang & Zhao, 2007). The presence of b-(1 / 3) linkage breaks the uniform structure of oat beta-glucan,
allowing it to form viscous solution upon solubilization (Xu et al., 2013). The structure of wheat beta-glucan is quite
regular due to the predominance of trimeric unit that contributes to its high gelling ability (Li et al., 2011). Beta-glucan

FIGURE 20.5 Structures of Beta-glucan from different sources. Courtesy: Du, B., Meenu, M., Liu, H., & Xu, B. (2019). A concise review on the
molecular structure and function relationship of b-glucan. International Journal of Molecular Sciences, 20(16). https://doi.org/10.3390/ijms20164032.
348 Nutraceuticals and Health Care

derived from yeast and mushroom composed of b-(1 / 3) linkage with occasional b-(1 / 6) linkages, which has greater
biological activity, than cereal beta-glucan (b-(1 / 3) and b-(1 / 4)) (Manners et al., 1973). Fungal beta-glucan is a
mixture of linear b-(1 / 3) linkage with short branch chains connected through b-(1 / 6) (Han et al., 2008; Manners
et al., 1973; Nakashima et al., 2018) while bacterial beta-glucan has linear unbranched chain of b-(1 / 3)-D-glucan
backbone (McIntosh et al., 2005). Seaweed/algae beta-glucan is comprised of straight chain of b-(1 / 3)-D-glucan
with high levels of b-(1 / 6) branches (Teas, 1982).
Beta-glucans are divided into two categories, soluble and insoluble, depending upon the linkage, former is having
b-(1 / 3/1 / 6)-D-linked glucose and later consists of b-(1 / 3/1 / 4)-D-linked glucose. Based on the source and
extraction method, beta-glucan can exist in a range of conformations such as random coils, helices (single, double, or
triple), rodlike shape, and wormlike shapes. The molecular weight of beta-glucan varied from 102 to 106 Da, depending
upon the origin.
Each type of beta-glucan encompasses different molecular backbone and level and/or length of branching which
eventually affects its water solubility, ability to form aggregates, conformation (shape, size, and structure), molecular
weight, techno-functional properties, and its physiological impact (Bernstein et al., 2013; Kaur et al., 2019; Volman et al.,
2008). The structure of beta-glucans resembles the structure of cellulose but the only difference is being that in b-(1 / 3)-
glycosidic linkage which results in twist in the chain and gives stability to beta-glucan toward aggregation (Ahmad et al.,
2012). Many reports hold up this assumption that presence of b-(1 / 3)-glycosidic bond reduces the solubility and in-
creases the tendency to gel (Ahmad et al., 2012; Böhm & Kulicke, 1999; Cui et al., 2000). Two major oligomers, DP3 and
DP4, explain more than 90% structure of beta-glucan from cereals. The ratios of DP3 to DP4 in barley, oats, and wheat are
2.8e3.4, 2.1e2.4, and 3.0e3.8, respectively (Wood, 2011). Physical and rheological properties are mainly governed by
cellotriosyl units in beta-glucan. Therefore, it is necessary to identify the exact length and distribution of linear b-(1 / 4)-
D-glycosidic linkage area, which results in understanding the structure of beta-glucan for its practical utility.
The physiological activity of beta-glucan depends upon the conformational structure. For example, it was recorded by
Ross et al. (1999) that beta-glucans with beta-(1 / 3)-glycosidic linkage with beta-(1 / 6) linkage as side chain possess
strong immune action and have higher pro-inflammatory cytokine stimulation. The physiological effect is dependent on the
type of linkage, side change, and its solubility. It is proved that soluble glucans can induce inflammatory cytokine
stimulation strongly than less soluble beta-glucan (Soltanian et al., 2009).

20.5 Mechanism of action

Beta-glucan is a functional food ingredient, recognized for its prebiotic activity. Prebiotics are the food ingredient that
promotes the growth of beneficial microbiota in the gastrointestinal tract. Both soluble and insoluble beta-glucans when
taken orally escape digestion in stomach and later soluble beta-glucan fermented by colon microbiota into butyrates,
lactates, acetates, and propionates which results in maintaining gut health. This effect of beta-glucan is illustrated in
Fig. 20.6. Insoluble beta-glucan influences the gut transit rate and modulates the bowel movement to prevent constipation.

FIGURE 20.6 Immunomodulatory effect of beta-glucan.

 Beta-glucan Chapter | 20 349

Moreover, consumption of beta-glucan restricts the consumption of nutrients especially glucose which helps to maintain
blood glucose levels and can be beneficial for diabetic patients. Beta-glucan binds with excess bile acids and helps to
interact with several mutagens and carcinogenic compounds for their removal from the body.
Ingestion of beta-glucan stimulates the macrophage activity, T-cells, reticuloendothelial system, natural killer (NK)
cells, and alternative pathways. Upon binding various processes occur such as cellular pathway activation and direct re-
ceptor activation. These cause pathogen elimination from the body and produce antibodies which improve the immunity.
Beta-glucan activates macrophages and increases their number and size which further stimulates the release of immu-
noglobulin, cytokines, and interleukin that provides dense to the host against various pathogenic conditions (Fig. 20.6).
Additionally, macrophage activation is mediated via toll-like receptor 2 (TLR2) which signals the production of tumor
necrosis factor-a (TNF-a) through the nuclear factor-kB (NF-kB) pathway.
Beta-glucan is recognized by numerous receptors present on the cell membrane such as dendritic cells, monocytes,
macrophages, and NK cells. Among them, important receptors of beta-glucan are CR3 (CD11b/CD18) and Dectin-1,
additional receptors include Toll-2, lactosylceramides, and scavenger receptors. The Dectin-1 receptor determines the
specificity for beta-glucan from the surface of macrophages and neutrophils (Brown et al., 2002) resulting in a variety of
cellular responses like endocytosis, phagocytosis, and oxidative burst (Brown & Gordon, 2003; Gantner et al., 2003).
Together Dectin-1 and TLRs can induce the production of pro-inflammatory cytokines and chemokines including TNF-
a, MIP-2, and IL-12 in macrophages and dendritic cells (Gantner et al., 2003).
It was observed by many researchers that beta-glucan has strong pleiotropic effects on cytokine production and
antibody responses. Some studies reported activation of B cells by beta-glucan that secretes some pro-inflammatory
lymphokines like IL-8 (Ali et al., 2015). This secretion required direct involvement of various molecules like Dectin-1,
transcription factors NF-kB and AP-1, and mitogen-activated protein kinase.

20.6 Bioavailability
The study of Rice et al. (2005) on soluble glucan oral delivery and its gastrointestinal absorption found that the
bioavailability of three different soluble beta-glucans ranges from 0.5% to 4.9%. These researchers observed that among
these three beta-glucans, laminarin has highest bioavailability followed by scleroglucan. Rice et al. (2005) mentioned
that the glucan phosphate has 0.5% bioavailability. Nakamura et al. (2016) determined the bioavailability of resistant
glucan (RG) and hydrogenated resistant glucan (HRG) using rats and humans. The tests were performed with repeated
measures design within subjects. RG and HRG were compared with glucose and fructo-oligosaccharides which are
nondigestible for their effects on blood glucose and insulin and hydrogen excretion in rats and humans. Fermentability
based on breath hydrogen excretion was used for measuring the available energy. This study indicated that only small
amounts of oligosaccharides in RG and HRG were digested as well as slightly fermented whereas the large portion of
carbohydrates with large molecular weight were digested to small extent and underwent fermentation slightly in healthy
humans. The study concluded that the bioavailability of RG as well as HRG was very little in both humans and rats. This
is due to the limited mobility of major components of RG though the oligosaccharide contained in minor component of
purified RG, and RG was highly mobile through intestinal microbes.

20.7 Stability, safety, and toxicology

20.7.1 Stability
Beta-glucan is regarded as one of the important macromolecules for its health benefits. Many of the health benefits,
including cholesterol lowering ability, hypoglycemic, and laxative effects, of beta-glucan are attributed to its high viscosity
and molecular weight (Brummer et al., 2012; Regand et al., 2009; Tosh et al., 2010; Wolever et al., 2010). The viscosity
properties of beta-glucan are governed by its molecular weight distribution (Lan-Pidhainy et al., 2007; Wood, 2007).
Therefore, it becomes imperative to understand the stability of beta-glucan and effect of various processing conditions on
physicochemical characteristics, particularly viscosity and molecular weight, of beta-glucan. Many studies have reported
that various methods of processing like application of high-pressure-assisted homogenization (Kivelä et al., 2009; Tosh
et al., 2010), ultrasound treatment (Vårum et al., 1992), addition of ascorbic acid (Kivelä et al., 2009), extrusion (Tosh
et al., 2010), and irradiation (Sung et al., 2009) result in decreasing the molecular weight of beta-glucan.
350 Nutraceuticals and Health Care

20.7.1.1 Effect of milling and germination

Milling and pearling did not significantly affect the beta-glucan content of barley (KeShun et al., 2009). Sullivan et al.
(2010) had observed that pearling results in insignificant increase in beta-glucan of barley grains at lower degree. Their
study further reported that middlings produced from whole barley and pearled barley had almost equal level of beta-glucan.
Germination significantly decreases the beta-glucan content of barley. This may be due to enhanced activity of beta-glucan
degrading enzyme, beta-glucanase (Nielsen & Munck, 2003; Wang & Zhao, 2007).

20.7.1.2 Effect of cooking and baking

Processing methods like cooking and baking also affect the various physicochemical parameters like viscosity, extract-
ability, molecular weight, and solubility as well as amount of soluble, insoluble, and total beta-glucan. Baking process
decreases whereas cooking decreases the content of soluble beta-glucan (Johansson et al., 2007). Cooking and bulgur
making significantly reduce the beta-glucan content of barley due to leaching of beta-glucan into cooked water (Köksel
et al., 1999). Cavallero et al. (2002) reported that the bread baking process did not alter the total beta-glucan content
significantly. However, Andersson et al. (2004) mentioned that beta-glucan in barley or wheat flour used in bread making
may be degraded by the action of endogenous enzymes present. The amount of soluble beta-glucan increases on hy-
drothermal treatment along with enzyme application followed by sonication. This increase is reported to be due to
improved solubility of beta-glucan during these processing methods (Izydorczyk et al., 2000).

20.7.1.3 Effect of extrusion

Extrusion cooking decreases the content of beta-glucan. Increasing the feed moisture, one of the important extrusion
processing parameters affecting quality of extruded products, from 15% to 22.5% at constant extrusion temperature of
150 C, results in 8% reduction of beta-glucan. Extrusion performed at 170 temperature and 22.5% feed moisture
exhibited 10% reduction in beta-glucan (Huth et al., 2000). Extrusion temperature is also reported to improve the solubility
of beta-glucans (Singh et al., 2007). This increase in solubility of beta-glucan is due to its release from insoluble
component as a function of temperature, pressure, and shear generated by screw during extrusion (Sharma & Gujral, 2013).
Contrary, Fadel et al. (1988) reported that extrusion cooking results in insignificant increase in the total beta-glucan
content.

20.7.1.4 Effect of roasting

The thermal processing was found to impact the beta-glucan differently and it depended on timeetemperature combination
and the nature of heat (Jood & Kalra, 2001; Sharma et al., 2011). The beta-glucan is also affected by roasting process
which is one of the important unit operations in development of grain-based products having long shelf life, better sensory
parameters, and retention of nutritional value (Gujral et al., 2011; Sharma et al., 2011). Sharma et al. (2011) mentioned that
the total beta-glucan content remains unchanged as an effect of roasting, however, heat processing decreases the solubility
of beta-glucan due to short heating time. This decreased solubility may be attributed to the polymerization of beta-glucan
during heat processing.

20.7.1.5 Effect of freezing and storage

Cooking and frozen storage of beta-glucan reduces its solubility and molecular weight which results in reduced cholesterol
lowering ability of beta-glucan (Lovegrove et al., 2000). The beta-glucan solubility was found to be reduced during frozen
storage of oat bran muffins. This decrease in solubility reduced the physiological effects in lowering postprandial blood
glucose (Lan-Pidhainy et al., 2007). Therefore, Lan-Pidhainy et al. (2007) advised avoiding the frozen storage of beta-
glucan in order to maximize its physiological effects. The viscosity as well as solubility of beta-glucan isolated from
bread prepared with fortification of beta-glucans sourced from barley were affected by storage conditions, including
storage at 20 C (Moriartey et al., 2011). The solubility and viscosity of beta-glucan extracted from porridge and bread
incorporated with oat bran was affected by freezing and freeze-drying due to escape of water from hydrated beta-glucan as
well as protein and starch. Furthermore, it was observed that the viscosity of beta-glucan subjected to freeze-drying and
freezing process performed with liquid nitrogen did not alter significantly (Gamel et al., 2013). Harasym et al. (2015)
reported that freeze milling results in reduction of beta-glucan molecular weight.
 Beta-glucan Chapter | 20 351

20.7.2 Safety and toxicity

Many oral toxicity studies (Babícek et al., 2007; Jonker et al., 2010; Delaney, Carlson, Frazer, et al., 2003; Delaney,
Carlson, Zheng, et al., 2003) have been conducted for investigating the beta-glucan safety. Many of these studies (Delaney,
Carlson, Frazer, et al., 2003) have demonstrated that consumption of large level of concentrated beta-glucan did not cause
any detrimental consequences.
The study of Delaney et al. (2004) further indicated that the oral consumption of concentrated beta-glucan sourced from
barley is not clastogenic. The intake level of beta-glucan recommended by FDA for lowering blood cholesterol is 50 mg/kg
body weight/day (FDA, 2005). The study of Jonker et al. (2010) showed that administration of beta-glucan equivalent to a
concentration of 5.8e5.9 g/kg body weight/day in Wistar rats is not linked with occurrence of toxic symptoms. It was
found that administration of beta-glucan in concentrated form extracted from barley did not result in inflammatory or any
other kind of consequences in CD-1 mice (Delaney, Carlson, Zheng, et al., 2003).
Salecan, one of the glucans produced by microorganism Agrobacterium sp. ZX09, did not exhibit any adverse effects in
the subchronic and acute toxicity study conducted on male and female mice. The oral toxicity study conducted with single
dose indicated that the salecan in mice has LD50 value of above 3 g/kg for both male and female mice. The mice fed with
salecan at concentration of 5% or below level did not show any adverse effects in the subchronic study. The mice fed with
salecan and without salecan had similar clinical pathology parameters, gain in body weight, consumption of food, and
overall health. The treated animals did not show any concentration-dependent effects. Salecan has not proved toxic at the
tested level and displayed no-observed-adverse-effect level (NOAEL) at 14.48 g/kg/day in mice during a study conducted
on mice fed for 90 days (Xiu et al., 2011). Chen et al. (2011) reported that beta-glucan intake not caused substantial
changes in feed consumption, body weight, clinical observations, and organ weights as evidenced from the results of
subchronic toxicity studies conducted on rats fed on beta-glucan extracted from mushroom at a dose level of 0e2 g/kg/day
for 90 days. The mushroom sourced beta-glucans were found to have NOAEL of 2 g/kg/day, which was also the highest
dose tested.

20.8 Health benefits

Beta-glucans are widely acknowledged and popularly recognized owing to their numerous health promoting activities
including reduction in blood glucose, cholesterol, obesity, many cardiovascular diseases. Beta-glucans are also widely
recognized for their anticarcinogenic, antitumor, antiinflammatory, and antiviral activities. Some of the prominent health
benefits of beta-glucan are discussed in following sections.

20.8.1 Regulation of blood glucose and insulin level

Beta-glucans are extensively known for their regulation of blood glucose as well as insulin level which is advantageous for
inhibiting the occurrence of diabetes. The cereal beta-glucans, in particular, are identified for their ability to regulate
glucose level in blood and are used for prevention of diabetes. It has been observed that even very low amount of beta-
glucans sourced from oat can significantly decrease glycemia as well as insulin (Ekström et al., 2017). The advantageous
result of beta-glucan extracted from barley on postprandial glycemia has also observed (AbuMweis et al., 2016). Several
mechanisms have been proposed for this effect of beta-glucan on lowering the blood glucose levels. Many studies (Gamel
et al., 2014; Östman et al., 2006; Regand et al., 2011) have attributed the viscosity of beta-glucan as a major cause for
lowering glycemia, insulin, and also cholesterol levels. However, one of the studies (Cavallero et al., 2002) has indicated
the relationship between dose of cereal beta-glucan and reduction in plasma glucose level. It has also been proposed that
the beta-glucans interact with intestinal mucus and thereby regulate the postprandial blood glucose as well as insulin level
(Mackie et al., 2016). Nevertheless, it has been observed by data on blood glucose response that the reduction of post-
prandial glycemic response by beta-glucan is due to coil overlap (Rieder et al., 2017; Tosh, 2013) and hence coil overlap
could be another potential mechanism for the above-described health effect of beta-glucan. The beta-glucan particles which
are the component of cell wall and occur in network like native structure might produce complex matrix within the cell wall
by encapsulation with proteins and starch. This developed matrix could then decrease accessibility of enzyme resulting in
decrease in metabolism of starch, thus reduce postprandial glycemic response (Zhang et al., 2017). The beta-glucans,
owing to their high viscosity, are also likely to play a vital role as physical hindrance for absorption of glucose (Abbasi
et al., 2016). Glucans also stimulate rise of insulin discharge and weakening of insulin resistance because of the more
solution viscosity prominently restoring and progressing the pancreatic islet beta cells integrity and structures of tissues
352 Nutraceuticals and Health Care

(Liu et al., 2016). Further, it is found that the concentration of gut peptides like YY and ghrelin influences decrease in
glucose or insulin level. Gut peptides play essential role in glucose homeostasis by affecting gut hormones (Baldassano
et al., 2017; Pradhan et al., 2013).

20.8.2 Reduction of cholesterol level and prevention of coronary heart diseases

Cholesterol lowering effect of beta-glucans is mentioned by many studies. The risk of coronary heart diseases significantly
gets lowered due to reduction in cholesterol level (Anderson et al., 2010; Bae et al., 2009). The detailed mechanism behind
reduction in cholesterol level is yet to establish comprehensively; however, several hypotheses have been proposed for
possible action of beta-glucans in lowering the cholesterol level. One of the present and chief hypotheses states that the
cholesterol reducing effect of beta-glucan may be attributed to binding of bile salts with beta-glucans of high viscosity in
small intestine thereby limiting their reabsorption and enhanced excretion (Gunness et al., 2012, 2016). The reduction in
bile reabsorption may be due to decreased rate of bile mass transfer from layers of intestinal mucus (Mackie et al., 2016).
Limited reabsorption of bile salts leads to reduction in total cholesterol content of blood and cholesterol content of low-
density lipoprotein. This reduction in cholesterol is significantly influenced by type as well as amount of fibers in diet
(Ghaffarzadegan et al., 2018). It is also observed that the beta-glucans result in advantageous reprofiling of minor bile acids
(Gunness et al., 2012, 2016) and beta-glucans from hull-less barley decrease low-density lipoprotein cholesterol levels in
plasma by encouraging the excretion of fecal lipids (Tong et al., 2015). The biological activities of beta-glucan are
dependent on its intrinsic functionality as well as the physical form of beta-glucan in which the beta-glucan is carried
forward for metabolism (Grundy et al., 2017).

20.8.3 Lowering of fat for obesity

Consumption of high amount of fat and low amount of fibers is considered as responsible for the onset of obesity. As a
dietary fiber, beta-glucans from various plant sources are suggested to be important in prevention of obesity resulting from
consumption of food with high fat and some biochemical indicators linked with obesity, fatty liver, and size of adipocyte
(Hu et al., 2015). It has been shown by Aoe et al. (2017) that the consumption of elevated amount of beta-glucan from
barley significantly reduces visceral fat obesity. The mechanisms behind prevention of obesity as a function of dietary
fibers include: (a) delay in gastric emptying rate and prolonged satiation afterward having meal as a result of decreased
intake of energy and increased consistency of intestinal matter due to inclusion of dietary fibers in diet (Drzikova et al.,
2005), (b) central appetite and obesity is suppressed by increased production of acetates and butyrates through fermentation
as a result of effect of dietary fibers on colon microflora and other products (Frost et al., 2014; Lin et al., 2012), and (c) the
food intake is reduced with the release of anorectic gut hormones PYY and GLP-1, which might promote increase in fatty
acids with short-chain length in the large intestine due to action of dietary fibers (Lin et al., 2012).

20.8.4 Reduction of blood pressure

Beta-glucan possesses the ability to lower the blood pressure. The viscosity properties of beta-glucan are supposedly the
mechanism of action for blood pressure lowering effect (Bai et al., 2019). However, many mechanisms have been claimed
for this action of beta-glucan. The viscosity effect of beta-glucan prompts reduction in the weight, which is proposed to be
linked with blood pressure lowering effect (Charlton et al., 2012; Chutkan et al., 2012; Khan et al., 2018). The viscosity of
beta-glucans is suggested to affect insulin resistance (Wong & Jenkins, 2007), which is a chief indicator of hypertension
(Montero, 2013). The renal sodium reabsorption and transmembrane transportation of ions, which reduces blood pressure,
is affected by high intake of viscous fibers (Schulman & Zhou, 2009; Zhou et al., 2010). This suggests that there is possible
association between blood pressure, insulin level, body fat, serum cholesterol level, and blood glucose. These various
aspects in human are likely to closely linked with each other and therefore even minor alteration in one aspect can affect all
other related aspects. All of these aspects are suggested to be affected by viscosity of beta-glucans, fermentation induced
formation of acetates and butyrates, and gut peptides (Bai et al., 2019).

20.8.5 Improving gut flora and laxative effect

The resident microbiota governs fate of beta-glucans at molecular level in human. The composition of human gut
microbiota is primarily driven by the utilization of complex polysaccharides by microorganisms. The syntenic mixed
linkage beta-(1,3)/beta-(1,4)-glucan utilization locus (MLGUL) works as genetic indicator for catabolism of MLG (Tamura
et al., 2017).
 Beta-glucan Chapter | 20 353

The mechanism behind laxative effect of beta-glucans is attributed to gut microbiotaeaccelerated intestinal peristalsis.
Moreover, it is suggested that there are two ways in which the mechanisms that are responsible for laxative effect work.
First being the stimulation of water and mucous secretion in large intestine by mechanical irritation of gut mucosa with
large or coarse insoluble fiber particles. Second being resistance to dehydration due to capacity of gel-forming soluble
fibers to hold more water. However, the above-described both mechanisms need fiber to not undergo fermentation process
and stay unchanged during passage through the large intestine for which it is necessary that the fibers must be available in
stool. Furthermore, these two mechanisms effect higher water amount in stool producing soft, bulky, and easily passed
stools (McRorie & McKeown, 2017).

20.8.6 Antitumor effect

The potential of beta-glucans as antitumor agent has long been recognized. Mo et al. (2017) reported that the beta-glucans
extracted from yeast exert antitumor effects. Beta-glucans from Diaporthe sp. endophytes display antiproliferative action
over carcinoma of human breast (MCF-7) and cells of hepatocellular carcinoma (Orlandelli et al., 2017). The beta-glucan,
lentinan, produced by L. edodes was capable of inducing apoptosis of S180 cells mediated through mitochondrial pathways
(Zhang et al., 2017). The beta-glucan prompted strong expression of caspase-12 in Me45 and A431 cancer cell lines might
be responsible for the strong antitumor properties demonstrated by low-molecular-weight beta-glucan of oat (Choromanska
et al., 2015). Avena sativa sourced beta-glucan is reported to have proapoptotic properties over human melanoma HTB-140
cells in vitro. The induction of apoptosis by beta-glucans from oat is due to concentration-dependent rise in activation of
caspase-3/7 and appearance of phosphatidylserine on outer surface of cellular membrane, where it binds to annexin
V-FTIC. Further, the mitochondrial pathway for apoptosis is evidenced from decrease in level of intracellular ATP with
decreasing mitochondrial potential. Increase in quantity of apoptotic cells and cells in G1 phase as well as reduction in
quantity of cells in G2/M is evidenced from analysis of cell cycle (Parzonko et al., 2015). The repression of genes related to
the G1 phase of the cell cycle may be attributed to antiproliferative activity of beta-glucan and to association of beta-glucan
with CCR5 receptor (Malini et al., 2015).
Beta-glucans isolated from S. cerevisiae also exert antitumor effect, which may be due to polysaccharides having
immune-stimulating properties their apoptosis inducing features (Mo et al., 2017). Beta-glucan has reported to retard
legumain action due to internalization of legumain into macrophages, which is induced by dectin-1 receptor. It was also
observed that in RAW 264.7 macrophages, the tumor necrosis factor-alpha inducing activity of particulate beta-glucans is
more compared to partially water-soluble glucans and water-soluble glucans (Berven et al., 2015).
Though the mechanisms associated with antiproliferative action of beta-glucan are yet to be comprehensively un-
derstood, it seems that the inhibitory activity necessitates presence of copyranose primary chain which is substituted at O-6
position by glucopyranose, since the branches may have vital contribution in cell death induction by favoring increased
efficacy of interaction with cell receptors located in tumor cells (Pires et al., 2017; Ruthes et al., 2015).

20.8.7 Antiinflammatory action

The beta-glucans delivered orally exhibit aggravated inflammation of intestine induced by means of dextran sulfate sodium
at mucosal level (Heinsbroek et al., 2015); however, beta-glucans are believed to possess positive antiinflammatory
activity. Yeast originated beta-glucans are reported to recover alterations in DSS prompted lesions with mucosal
inflammation and intestinal hindrance. This recovery effect is believed to be achieved through hindering the expression of
mediators of inflammatory nature and improving the expression of proteins with tight junction linked to permeability of
intestines (Han et al., 2008; Manners et al., 1973; Nakashima et al., 2018). The low-molecular-weight beta-glucans from
oat reduce the enteritis group in rats as a result of elevated defense to oxidation (Suchecka et al., 2015). The
low-molecular-weight beta-glucans activate leukocytes through stimulation of nuclear transcription factors whereas the
high-molecular-weight beta-glucans stimulate leukocytes and govern the synthesis of pro-inflammatory cytokines and
chemokines (Brown & Gordon, 2003). The beta-glucans extracted from yeast improve gene expression of Cath-2, AvBD-4,
and AvBD-10 during initial stage of infection and gene expression of Cath-1, Cath-2, and AvBD-1 during progressive stage
of infection in the gut. These beta-glucans, however, reduce the expression of AvBD-10 and LEAP-2 mRNA level during
progressive stage of infection. This increase and decrease of abovementioned gene expression is reported to improve
necrotic enteritis induced by perfringens (Tian et al., 2016). Though the beta-glucans are regarded as antiinflammatory
agents, their role in whole immune system remains unclear because some hypotheses describing relationships among
various proposed mechanisms of action have not been confirmed (Bai et al., 2019).
354 Nutraceuticals and Health Care

20.9 Conclusion
Beta-glucan is one of the widely recognized polysaccharide for its therapeutic as well as functional properties. Beta-glucan
naturally occurs in many foods, including cereal grains, particularly oat and barley. Beta-glucan can also be sourced from
some microorganisms like yeasts. Beta-glucan shows structural variability based on its source. Limited studies are con-
ducted on bioavailability of beta-glucan. The bioavailability of soluble beta-glucan is considerably less. Technical in-
terventions may be applied on improving the beta-glucan bioavailability. Beta-glucan shows high stability during many
conventional methods of processing. Effects of nonthermal processing on beta-glucan need to be undertaken. Oral
administration of beta-glucan in animals is safe and has no toxic effects at high doses. The safety and toxicity studies may
be conducted at even increased levels of beta-glucan which can lead to increasing the incorporation of beta-glucans in
higher amounts in the processed foods. Many mechanisms of actions are proposed for the health benefits of beta-glucan;
however, some mechanisms are yet to be fully established and comprehended.

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