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A review of β-glucans as a growth
promoter and antibiotic alternative
against enteric pathogens in poultry
M.I. ANWAR
1
*, F. MUHAMMAD
2
, M.M. AWAIS
1
and M. AKHTAR
1
1
Department of Pathobiology, Faculty of Veterinary Sciences, Bahauddin Zakariya
University, Multan, Pakistan;
2
Institute of Pharmacy, Physiology and
Pharmacology, University of Agriculture, Faisalabad, Pakistan
*Corresponding author: drirfananwar@hotmail.com
The emergence of microbial challenges in commercial poultry farming causes
significant economic losses. Vaccination is effective in preventing diseases of
single aetiology while antibiotics have an advantage over vaccination in
controlling diseases of multiple aetiologies. As the occurrence of antibiotic
resistance is a serious problem, there is increased pressure on producers to
reduce antibiotic use in poultry production. Therefore, it is essential to use
alternative substances to cope with microbial challenges in commercial poultry
farming. This review will focus on the role of β-glucans originating from yeast
cell wall (YCW) as a growth promoter and antibiotic alternative. β-glucans have
the ability to modulate the intestinal morphology by increasing the number of goblet
cells, mucin expression and cells expressing secretory IgA (sIgA) with increased
sIgA in the intestinal lumen and decreased bacterial translocation to different
organs. β-glucans also increase the gene expression of tight junction (TJ) proteins
which maintain the integrity of the intestinal wall in broiler chickens. However,
further studies are required to optimise the dosage and source of β-glucans to
determine effects on growth performance and mechanisms against enteric
pathogens.
Keywords: antibiotic alternative; YCW β-glucans; enteric pathogens; poultry
Introduction
The poultry industry is one of the largest and rapidly growing industries worldwide and
the prevalence of pathogenic microbes challenges practical management and causes
significant economic losses. Additionally, the downstream industry, including
slaughter, shipping and retail marketing tends to suffer from carcasses contamination
by pathogens (Mainali et al., 2014). Vaccination can prevent diseases of a single
aetiology. Antibiotics have some benefits over vaccination when multiple infections
© World's Poultry Science Association 2017
World's Poultry Science Journal, Vol. 73, September 2017
Received for publication September 10, 2016
Accepted for publication February 21, 2017 651
doi:10.1017/S0043933917000241
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0043933917000241
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and environmental stresses produce diseases. Since most antibiotics produced are used in
diets of livestock animals, the increased occurrence of antibiotic resistance in microbes
further limits the use of antibiotics in poultry.
Contemporary international legislation has withdrawn antibiotics in poultry production
as a growth promoter. Consequently, it is necessary to find alternates for microbial
challenges in domestic fowl. Certain research attention has concentrated on the
possible use of YCW β-glucans in chickens to combat the limitations mentioned
above. Subsequent studies have demonstrated that dietary supplementation of β-
glucans can reduce the severity of enteric pathogen infection (Huff et al., 2006; Shao
et al., 2013), increase phagocytosis by macrophages after bacterial infection (Chen et al.,
2008) and improve growth performance (Cho et al., 2013). Based on the characteristics
of β-glucans, this review will focus on their effects as growth promoters and antibiotic
alternatives against enteric pathogens in poultry.
Yeast cell wall β-glucans
Saccharomyces cerevisiae, is a unicellular yeast that has been developed to be used in
animal feeds (Hooge, 2004; Rosen, 2007). Broiler production can be improved by
maintaining the intestinal integrity with the inclusion of yeast in their diets (Santin et
al., 2001; Zhang et al., 2005; Baurhoo et al., 2007). Typically, the yeast cell wall (YCW)
is composed of β-glucans, mannoproteins, and chitin with varying degrees of
polymerisation, as presented in Table 1.
Table 1 YCW components with their molecular weight and degree of polymerisation (DP).
Macromolecule Cell wall mass Mean molecular weight,
(%, dry weight)
a
kDa (DP)
Mannoproteins 35–40 Highly variable
1,3 β-Glucan 50–55 240 (1500)
1,6 β-Glucan 5–10 24 (150)
Chitin 2 25 (120)
a
=Klis et al., 2002; Freimund et al., 2003; Lessage and Bussey, 2006; Kwiatkowski et al., 2009.
YCW components, presented in Figure 1, can vary depending upon growth conditions,
time of harvesting (Klis et al., 2002; Aguilar-Uscanga and Francois 2003; Klis et al.,
2006) and most importantly, strains of yeast (Hahn-Hägerdal et al., 2005). In general,
baker's yeast is used for producing high quality β-glucans with good therapeutic
applications (Kim et al., 2007). The processes involved in the separation of β-glucans
have been heavily patented., and there are various methods for separation of β-glucans
from yeasts (Borchani et al., 2016).
β-glucans as a YCW component, are composed with polymerisation of glucose through
β-1,3/1,6 glycosidic linkages. It has the ability to bind with numerous types of cell
surface receptors on macrophages and polymorphonuclear cells (PMNCs) by binding
to Dectin-1 or CR3 receptors (Palićet al., 2006) with subsequent activation of
macrophages/lymphocyte, production of inflammatory cytokines, leukotriene,
interleukins and stimulation of phagocytosis for microbial killing (Brown and Gordon,
2003; Brown et al., 2003). Figure 2 demonstrates activation of macrophages by β-
glucans binding to Dectin-1 receptor.
652 World's Poultry Science Journal, Vol. 73, September 2017
β-glucans as antibiotic alternative: M.I. Anwar et al.
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Figure 1 Structure of yeast cell wall.
Figure 2 Activation of macrophages by YCW β-glucan binding to Dectin-1 receptor.
Furthermore, YCW β-glucans can modulate the function of neutrophils that assists in
disease resistance. Benefits of YCW β-glucans in poultry production can be partially
attributed to enhanced proliferation and phagocytosis by avian macrophages and
heterophils (Guo et al., 2003; Lowry et al., 2005). Broilers fed diets with YCW β-
glucans supplementation showed amplified cell mediated (Chen et al., 2003; Chae et al.,
2006) and humoral immune responses (Guo et al., 2003; Zhang et al., 2008). This
immune enhancing ability can remove Salmonella enterica and Escherichia coli,
economically important pathogens, from the digestive tract (Lowry et al., 2005; Huff
et al., 2010). Furthermore, YCW β-glucans can enhance broiler performance under
unhygienic conditions through increases in villi height, uniformity and integrity,
leading to improved intestinal function and gut health (Huff et al., 2006; Yang et al.,
2007). The YCW β-glucans from different sources have large variations in their structure
that ultimately affect their physiological functions (Volman et al., 2008).
World's Poultry Science Journal, Vol. 73, September 2017 653
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Alternative to antibiotic growth promoters
Feeds containing no chemical additives are increasingly used in poultry nutrition. Since
antibiotic growth promoters have been discredited by consumer associations, there has
been an increasing trend in formulating diets without supplementation of synthetic drugs
and chemical additives and many scientists are dedicated in finding natural alternatives to
replace antibiotics as growth promoters (Langhout, 2000; Mellor, 2000).
There are numerous studies on the role of YCW β-glucans as growth promoters in
poultry birds. A growth improving effect of YCW β-glucans at 0.1% dietary
supplementation in an unchallenged setting was observed (Cho et al., 2013), which
showed improved body weight gain in broilers compared to control groups. In
another study, Cox et al. (2010b) noted that dietary supplementation with 0.02 or
0.1% β-glucans had no significant differences on growth performance of broilers with
or without an Eimeria challenge. However, Zhang et al. (2008) reported a growth
improving effect of β-glucan supplemented at 50 and 75 mg/kg in broilers. Rathgeber
et al. (2008) found that 40 ppm yeast β-glucans could increase body weight in broilers
during the grower phase, and an increase in relative weight of the gizzard was observed,
enhancing the digestive capacity of broilers which may account for the growth effect.
Broiler chickens supplemented with 22 ppm β-glucans in feed showed some
improvement when infected by E. coli (Huff et al., 2006). In another study, the
progressive effects of β-glucans were noted in birds reared on litter pen than those in
the cages (Chae et al., 2006). The variations in the results reported could be due to
reasons including differences in the sources of β-glucans (species and strains), purity,
composition, dosage or presence/absence and type of pathogen challenge (Zhang et al.,
2008; Cox et al., 2010b). More studies to optimise the dosage and source of β-glucans for
consistent results in broiler production are required.
β-glucans originating from YCW have different effects on the visceral organs. A study
by Cho et al. (2013) showed that 0.1% dietary supplementation of YCW β-glucans in an
unchallenged trial resulted in an improvement in the relative weight of the spleen and the
bursa of Fabricius. The effects of β-glucans on the relative weight of the spleen and the
bursa of Fabricius varied with studies, depending on the dose and the presence of
pathogen challenges (Rathgeber et al., 2008; Cox et al., 2010a; Salarmoini and
Fooladi, 2011). In another study broilers supplemented with 40-50 mg/kg of β-
glucans in feed showed an increase in the relative weight of the thymus, bursa of
Fabricius and spleen (Guo et al., 2003; Zhang et al., 2008). Chickens supplemented
with mannoprotein and β-glucans showed improved thymus weight (Morales-Lopez et
al., 2009), and β-glucan supplementation improved red blood cells count (Guo et al.,
2003) and increased the villus height of the jejunum mucosa of chickens (Morales-Lopez
et al., 2009), and subsequently may increase the surface area for nutrient absorption.
Further studies are required to delineate the mechanisms of the effect of β-glucans on
primary immune organs and other visceral organs weight in relation to growth
performance.
Activity against enteric pathogens
In humans, food-borne gastroenteritis is mainly caused by Salmonella enterica serovar
typhimurium from contaminated poultry products (Zhang et al., 2003; Liljebjelke et al.,
2005). Severe colonisation by Salmonella typhimurium in the gastrointestinal tract can
damage intestinal barriers and may cause bacteremia in both domestic fowls and humans
(Jepson et al., 2000; Zhang et al., 2003; Griffin and McSorley, 2011). Moreover,
654 World's Poultry Science Journal, Vol. 73, September 2017
β-glucans as antibiotic alternative: M.I. Anwar et al.
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Salmonella infection causes high mortality and decreases the production of poultry birds
(Fasina et al., 2008). Colibacillosis, caused by E. coli, is another important enteric
disease in domestic fowl and causes air sacculitis, decreased BW gain, and increased
faecal and carcass contamination (Barnes et al., 2003; Russell, 2003). Gastrointestinal
parasites constitute major stress factors which reduce growth performance and nutrient
utilisation in broilers. Coccidiosis by an intracellular protozoan belonging to the genus
Eimeria (Dalloul and Lillehoj, 2005) can disrupt the intestinal lining resulting in gross
lesions and decreased growth performance by malabsorption of nutrients (Brake et al.,
1997). Controlling these infections is needed for food safety, as well as for both animal
and human health.
Various prophylactic approaches including antibiotics, vaccines, genetic selection for
improved immunity and standard hygienic conditions, are being used to eradicate these
infections from poultry birds (Vandeplas et al., 2010). Dietary supplementation with
yeast β-glucans may be beneficial in controlling these challenges.
Defence mechanisms in the gastrointestinal tract (GIT)
In addition to nutrient absorption, the lining of the digestive tract acts as a barrier against
pathogen invasion. Goblet cells secret mucins to attract microbe adhesion to increase
elimination of microbes, and thus prevent physical and chemical injury by pathogens
(Deplancke and Gaskins, 2001; Iijima et al., 2001). When the mucus layer is disturbed,
adhesion of microbes to the intestinal epithelial surface may increase epithelial
permeability and reduce the absorption of nutrients. Intestinal secretory
immunoglobulin A (sIgA) is a line of defence in protecting the intestinal epithelium
from pathogenic microorganisms and maintaining the mucosal layer (Mantis et al., 2011).
Intestinal integrity is maintained by tight junctions (TJ) proteins including claudins,
junctional adhesion molecules and occludin (Tsukita et al., 2001; Schneeberger,
2004). Claudin and occludin families thus maintain intestinal integrity to form a solid
barrier from pathogen invasion (Fanning et al., 1998) as presented in Figure 3 (a).
Damage in GIT due to enteric pathogens
Invasion of enteric pathogens such as Salmonella typhimurium causes loss of integrity
and dysfunction of the intestinal epithelium (Clark et al., 1998; Sears, 2000). Salmonella
typhimurium infection can down regulate TJ protein expression in the intestinal epithelial
cells, damage intestinal barrier function, and thus facilitate bacterial invasion into the
blood stream (Jepson et al., 2000; Kohler et al., 2007). Broilers infected by Salmonella
spp. have been reported to have decreased villi height in the jejunum and altered ratio to
crypt depth, a reduced number of goblet cells with increased sIgA production and its
expressing cells (Beal et al., 2004; Withanage et al., 2005; Rahimi et al., 2009;
Revolledo et al., 2009; Fasina et al., 2010; Marcq et al., 2010; Shao et al., 2013) as
presented in Figure 3(b). Cytokines, including interleukin-6 (IL-6), IL-1β, IL-8 and
interferon-γ, play a pivotal role in the regulation of intestinal TJ barrier with
Salmonella spp. infection (Withanage et al., 2004; Withanage et al., 2005; Fasina et
al., 2008; Al-Sadi, 2009; Groschwitz and Hogan, 2009; Turner, 2009). TJ proteins
including occluding, claudin-1 and claudin-4 expression declined in the jejunum
during Salmonella spp. infection in broilers (Zhang et al., 2012). Decrease of TJ
protein expressions increases epithelial permeability, interrupts intestinal barriers and
allows leakage and invasion of antigens, pathogens and bacterial toxins into the
World's Poultry Science Journal, Vol. 73, September 2017 655
β-glucans as antibiotic alternative: M.I. Anwar et al.
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circulation (Kucharzik et al., 2001; Tsukita et al., 2001; Schneeberger, 2004). Increased
intestinal permeability has been shown to increase the infiltration of pathogenic bacteria
into distinct organs including the liver and spleen (Kops et al., 1996; Jepson et al., 2000;
Kohler et al., 2007; Revolledo et al., 2009).
Mechanisms by which β-glucans act as an antibiotic alternative
Yeast β-glucans have been shown to have positive effects against various infectious
diseases by improving immunity, while its role as an antibiotic alternative in domestic
fowl against enteric pathogens is not yet completely explored. β-glucans have various
mechanisms to improve the growth performance of domestic fowls, in which dietary
supplementation has been shown to increase the intestinal clearance of numerous
imperative pathogens such as E. coli and Salmonella spp. (Lowry et al., 2005; Huff
et al., 2010) by protecting intestinal barriers, stimulating phagocytosis and suppressing
pathogen invasion into organs (Lowry et al., 2005). Previous studies have suggested that
β-glucan supplementation can enhance goblet cell number and villus height in the ileum
(de Los Santos et al., 2007; Morales-Lopez et al., 2009), as well as restore villus damage/
loss by Salmonella spp. challenge (Shao et al., 2013). In addition, β-glucans
supplemented at 0.1% level of the diet improved villus height and crypt depth and
their ratio in the duodenum and ileum, upregulated mucin-2 production from goblet
cells, modulated intestinal profile of cytokines, and reduced lesion severity and
inflammation in the intestine of broilers during Eimeria/coccidia infection (Cox et al.,
2010a; 2010b). Another mechanism by which YCW β-glucans can act as an antibiotic
alternative is attributed to increased sIgA secretion and numbers of goblet cells that
maintain the integrity of the mucous protective layers for expulsion of enteric
pathogen invasion (Brandtzaeg, 2010). β-glucan supplementation has been shown to
increase jejunal TJ protein expressions such as occluding and claudin-1, leading to the
increased removal of Salmonella Typhimurium (Shao et al., 2013) as illustrated in Figure
3 (c), while it also inhibits the colonisation and invasion of Salmonella in the caeca and
liver, respectively (Lowry et al., 2005; Chen et al., 2008; Revolledo et al., 2009).
These mechanisms of yeast β-glucans relate to the improvement of the intestinal
epithelial cell functions but no mechanism regarding direct killing or binding of β-
glucans with enteric pathogens is available to date. However, various studies relating
to microbial killing by activation of macrophages/lymphocytes are available.
Conclusions
YCW β-glucans can act as a safeguard in domestic fowl against enteric pathogens by
increasing sIgA secretion, goblet cell numbers, upregulating expression of intestinal TJ
proteins and cytokines. Moreover YCW β-glucans merits being considered as a growth
promoter in broilers. As a growth promoter and antibiotic alternative, further studies are
required to optimise the dosage and source of β-glucans to determine its effects on
growth performance and mechanisms against enteric pathogens. In conclusion, β-
glucans may be used as a growth promoter and antibiotic alternative.
656 World's Poultry Science Journal, Vol. 73, September 2017
β-glucans as antibiotic alternative: M.I. Anwar et al.
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0043933917000241
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Figure 3 Intestinal morphology. (a) normal goblet cells, TJ proteins, sIgA, mucous, intestinal villi, B and
T lymphocytes, plasma and dendritic cells, (b) disruption of TJ proteins, reduced villus height, decreased
goblet cells, mucous, increased sIgA, dendritic cells, plasma cells, T and B lymphocytes by enteric
pathogens, (c) increased goblet cells, mucous, sIgA, T and B lymphocytes, plasma cells, and dendritic cells
with β-glucans and reduced enteric pathogens.
Conflict of interest statement
None of the authors of this paper has a financial or personal relationship with other
people or organisations that could inappropriately influence or bias the content of the
paper.
World's Poultry Science Journal, Vol. 73, September 2017 657
β-glucans as antibiotic alternative: M.I. Anwar et al.
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0043933917000241
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