Xylooligosaccharides As Prebiotics From Agricultural By-Products

Bioactive Carbohydrates and Dietary Fibre 5 (2015) 62 71
Available online at www.sciencedirect.com
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Xylooligosaccharides as prebiotics from agricultural

by-products: Production and applications
A.K. Samantan, Natasha Jayapal, C. Jayaram, Sohini Roy, A.P. Kolte,
S. Senani, Manpal Sridhar
Feed Additives and Nutraceuticals Laboratory, National Institute of Animal Nutrition and Physiology, Adugodi, Hosur
Road, Bangalore-560030, Karnataka, India
ar t ic l e in f o
abs tra ct
Article history:
The growing demands of novel food products for well-being and age related issues coupled
Received 15 October 2014
with increasing health care expenditure has attracted global attention on prebiotics.
Received in revised form
During the end of twentieth century, a great concern was expressed for transfer of
23 December 2014
antibiotic resistance genes from animal to human through food chains leading to the
Accepted 24 December 2014
concept of preventive medication. Xylooligosaccharides (XOS) is the only nutraceutical

that can be produced from lignocellulosic biomass. Production of XOS from agricultural
Keywords:
residues offers great scope to the nutraceutical industries as the raw material is cheap and
Prebiotics
abundantly available. The major advantages of XOS consumption, apart from selective
Xylooligosaccharides
growth stimulation of benecial gut microora, include reduction of blood glucose and
Agricultural residues
cholesterol, reduced pro-carcinogenic enzymes in gastrointestinal tract, enhanced mineral
Gut health
absorption from large intestine and immune-stimulation. The sweet taste of XOS enables it
to be used as an articial sweetener. Owing to the ban on antibiotics as a feed supplement
in livestock, the XOS could be future alternatives to guard gastrointestinal tract from the
onslaught of pathogenic microora. This will sustain the productivity and improve the
quality of animal products to full the demands of value added livestock products.
Therefore, XOS could be the future sought after molecules in preventive medicine.
& 2014 Elsevier Ltd. All rights reserved.
Contents
1.
2.
3.
4.
5.
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Prebiotics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Structural properties of xylooligosaccharides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Agricultural residues for xylan fractionation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Xylooligosaccharides production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
5.1. Chemical process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
5.2. Enzymatic process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
5.3. Autohydrolysis process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Corresponding author. Tel.: 91 80 25711304x313; Mobile: 91 9449447397; fax: 918025711420.

E-mail address: drashiskumarsamanta@gmail.com (A.K. Samanta).
http://dx.doi.org/10.1016/j.bcdf.2014.12.003
2212-6198/& 2014 Elsevier Ltd. All rights reserved.
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6.
Biological role of xylooligosaccharides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

6.1. Selective growth stimulation of gut microora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6.2. Antioxidant property of XOS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6.3. Short chain fatty acid production. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6.4. Immune stimulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6.5. Lipidaemic effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
7. Safety aspects of XOS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
8. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1.
Introduction
In the light of increased health consciousness and consumer

awareness in todays world, everyone prefers naturally occurring bioactive molecules in place of modern therapeutic agents
for their health and well-being. These bioactive compounds
work on the principle of Prevention is better than cure and
exhibit the benets by enhancing quality and expectancy of
human life. The great Greek physician Hippocrates (460370
BC) advocated Let food be thy medicine and medicine be thy
food. With respect to the above principle of Hippocrates, in
todays world, consumers also prefer foods with additional
attributes along with routine nutritional qualities. In this
context, although several biologically active compounds are
being tested; prebiotics seem to be the preferred one as it
demonstrates benecial effects over several physiological
functions such as selective growth stimulation of benecial
gut microora, enhanced mineral absorption, cholesterol lowering, glucose homeostasis, pathogen exclusion, immune
modulation, antioxidant, anti-carcinogenic properties, etc.
Although, oligosaccharides are available as low calorie bulking
agents since eighties, but oligosaccharides based prebiotics
have garnered much interest recently due to discovery of
multidimensional benecial roles on human lives and livestock. Among the prebiotic compounds, xylooligosaccharides
(XOS) appears to be a promising one, since these can be
sourced from agricultural crop residues that are inexpensive,
abundant and renewable in nature.
2.
Prebiotics
At the dawn of twenty rst century, everybody is concerned

about contaminating gut pathogens and seeking remedies to
avoid ingestion and subsequent ill effects. Researchers are
entrusted with the task to supply selective nutrients to stimulate the growth and multiplication of a class of gut microora
that are known to play several benecial roles in both humans
and domesticated animals like poultry, dog, swine, etc. Evidently, this culminates into the emergence of a new concept in
the history of functional food science; popularly known as
prebiotic, which came into light in the year of 1995. Prebiotics
may be dened as Non-digestible food ingredients that benecially affect the host by selectively stimulating the growth
and/or activity of one or a limited number of bacteria in the
colon (Gibson & Roberfroid, 1995). As some of its features
match with dietary bre, its denition passes through several
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steps of modications during subsequent period. Considering

the importance of prebiotics on overall human health, the Food
and Agricultural Organization (FAO, 2007) constituted a technical committee to revisit the denition of prebiotic. According to
FAO, A prebiotic is a non-viable food component that confers
health benet on the host associated with modulation of
microbiota. FAO has suggested three criteria for qualifying a
substrate as prebiotic:
(1) Component: Not an organism or drug; a substance that
can be characterized chemically and in most cases these
will be of food grade.
(2) Health benets: Measurable and not due to absorption of
the component in the blood or due to the component
acting alone.
(3) Modulation: Deliver changes in the composition or activities of microbiota of the target host including fermentation, receptor blockage or others.
Therefore, a prebiotic can be a bre but all bres are not

prebiotic. Subsequently, attempts were made to broaden the
site of action of prebiotics such as skin, oral cavity and female
genital tract in addition to its fundamental action site i.e.
gastrointestinal tract (Pineiro et al., 2008). Recently, the proposers of prebiotic concept have opposed the philosophy of
broadening its application sites such as vagina and skin and
wanted to restrict once again to the gastrointestinal microora.
Therefore, the accepted denition remains closer to the 1995
concept: A prebiotic is a selectively fermented ingredient that
results in specic changes in the composition and/or activity of
the gastrointestinal microbiota thus conferring benet(s) upon
host health (Gibson et al., 2010). In line with this principle,
Roberfroid et al. (2010) suggested that the concept of prebiotic
should stand with the following principal theory: The selective
stimulation of growth and or activities of one or a limited
number of microbial genus(era)/species in the gut microbiota
that confer(s) health benets to the host. Accordingly, the
prebiotics concept is crowned with selective stimulation of
growth and activity of single or multiple gut microora in order
to demonstrate quantiable health benets following its consumption. However, irrespective of the changes in dening the
concept with passage of time, an ideal prebiotic should have
the following criteria (Samanta et al., 2007):
Selective fermentation by benecial gut microora (evident from in vitro or in vivo experiments).
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Modulation of gut microora homeostasis towards bene

cial sides resulting into increase of population or metabolic activities.

Ensures host health and well-being (enhances productivity or product quality in animals).
Originated from plant or synthesized by microorganism or
their enzymes.
Maintains structural integrity while passing through different parts of the gastrointestinal tract.
No residue problems.
Ideally to be used as a food/feed additive.
Compatibility with other food/feed ingredients.
Amongst the prebiotics, XOS occupies a signicant niche

because of their multi-dimensional inuences on human
health in addition to their potentiality to work against several
gastrointestinal disorders. Further, it is thought to be important
as its production basically relies on lignocellulosic materials,
which are unsuitable for human consumption and available in
plenty all over the world. XOS is reported to be present naturally
in honey, bamboo shoot, fruits, vegetables etc.; but its concentration is inadequate to exhibit prebiotic effects. In order to
ensure global food security, every step towards inching up the
productivity of crops will obviously generate additional quantity
of lignocellulosic biomass that will be a major concern to

farmers, industrialists, environmentalist and civic authorities.
Thus, the mechanism to translate the biomolecule of overproduced lignocellulosic biomass into value added products
such as XOS could really pose to be a boon to all concerned
authorities as well as for improvement of both human and
livestock health.
3.
Structural properties of
xylooligosaccharides
XOS are sugar oligomers comprised of xylose units through
-(1-4)-xylosidic linkages viz.; xylobiose (2 monomers), xylotriose (3 monomers), xylotetrose (4 monomers), xylopentose (5
monomers), xylohexose (6 monomers) and so on (Kumar &
Satyanarayana, 2011). The common properties of XOS are
presented in Table 1. As the name implies, these are the
hydrolysis products of xylan; essentially present in most of
the lignocellulosic materials with varying degrees. The molecular formula of XOS is C5nH8n2O4n1; where, n 2 to 6. The
XOS derived from different raw materials may be present with
any one side chain such as acetyl, 4-O methyl derivative,
arabinofuranosyl and in such case it could be termed as
branched XOS (Aachary & Prapulla, 2008). The XOS remains
stable in acidic media and exhibits resistance to heat (Moura,
Table 1 Chemical and biological properties of xylooligosaccharides.

Property
Common name
Synonym
Molecular formula
Chemical family
Molecular weight
Physical status
Odour
Melting temperature
Decomposition temperature
Solubility
pH stability
Relative sweetness
Prole of sweetness
Cooling effect
Energy value
Carcinogenicity
Flavor enhancer
Humectants
Hygroscopicity
Absorbability at gastrointestinal
tract
Maillard reactions and
caramelization
Advantages of consumption
Recommended consumption dose

Side effects of excess dose
Regulatory status
Value/nature
Xylooligosaccharides
D-xylose-hexulose
C5nH8n2O4n1; n 2 to 6
Carbohydrate; xylobiose (DP 2), xylotriose (DP 3), xylotetrose (DP 4), xylopentose (DP 5) and xylohexose
(DP 6)
282 to 810 (X2 to X6)
Crystalline solid, colour depends on source xylan or purication or process of drying.
Nil
134 1C
120 1C
58% w/w at 21 1C
2 to 7
92% of sucrose when compared in 10% solutions
Emulates sucrose having faster onset like sucrose
Nil
1.5 kcal/g
Nil
Synergistic with high intensity sweeteners
Similar to sorbitol
Less than fructose
Malabsorption sugar in human
Browns similar to sucrose
Low calorie sugar alcohol having prebiotic effects
No elevation of blood glucose; suitable for diabetic patients
Antioxidant, cyto-protective
Offers scope for dietary supplements, benecial drug or drug adjuvant
812 g/day for healthy adult
Distension of intestine, nausea, atulence, diarrhoea etc
Excipient in drugs and non-foods: Generally Recognized as Safe (GRAS)
Excipient use in animal feeds: Generally Recognized as Safe (GRAS)
Application as antidiabetic drug: Under clinical trial
Gullon, Deminguez, & Parajo, 2006). XOS are non-carcinogenic

and regulate insulin secretion from pancreas in addition to
increase of mineral absorption from large intestine. It affects
bowel function through its mild laxative ability.
4.
Agricultural residues for xylan fractionation
As the name implies, the precursor for xylooligosaccharides

is xylan. Xylan is the polysaccharide accounting for 2535% of
the dry biomass of woody tissues of dicots and lignied
tissues of monocots and comprises up to 50% in some grasses
and tissues of cereal grains. The structure of xylan depends
on its source and fractionation process applied for its recovery. Most xylans are made up of main backbone of xylose
linked by -1,4-xylosidic linkages and substituted with arabinose, glucuronic acid, acetyl or methyl groups (Saha, 2003;
Samanta et al., 2012). Till today, several agricultural residues/
by-products (Table 2) have been evaluated to extract xylan by
different approaches. Among those, alkaline extraction
coupled with steam treatment resulted in recovery of more
than 90% of original xylan present in plant bio-materials
(Samanta et al., 2012, 2013).
5.
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could also be subjected to direct autohydrolysis under high

temperature and pressure to yield XOS.
5.1.
Chemical process
Albeit the prebiotic application of XOS appeared recently in the

eld of functional food science, the XOS production from the
lignocellulosic material was attempted as early as 1984 in order
to know the degradation pattern of oligomer and polymers of
Xylooligosaccharides production
In view of the rising demand of prebiotics around the world,

researchers are working for development of economic and
efcient process to produce XOS from diverse plant biomass
(Fig. 1). Following sculpturing of xylan from agricultural
biomass, XOS could be produced by chemical (Samanta
et al., 2012a), enzymatic (Jayapal et al., 2013) or combination
of chemical and enzymatic (Kokubo & Ikemizu, 2004) methods. Apart from this, the xylan rich agricultural plant biomass
Fig. 1 Schematic representation of xylooligosaccharides

(XOS) production from agricultural biomass.
Table 2 Extraction of xylan from various agricultural residues/ by-products.

Name of raw
materials
Extraction with
Yield
Reference
Cotton stalks
24% KOH1% NaHBH4
0.4 g/2 g of cotton stalks
Corn cobs
Corn stalks
12% NaOH with steam

10% NaOH 20 1C
83% of original xylan

54% of original sugars
Wheat straw
0.5 M NaOH at 55 1C
49.3% of original xylan
Poplar wood
Sugarcane
bagasse

Tobacco stalks
NaOH coupled with sonication

3% NaOH at 50 1C
12% NaOH coupled with steam
Two stage extraction with water and
1% NaOH
24% KOH1% NaHBH4
Akpinar, Ozlem, Kavas, Bakir, and Yilmaz

(2007)
Samanta et al. (2012a)
Ergues, Sanchez, Mondragon, and Labidi
(2012)
Ruzene, Silva, Vicente, Goncalves, and
Teixeira (2008)
Yuan, Xu, He, and Sun (2010)
Peng et al. (2009)
Jayapal et al. (2013)
Peng et al. (2010)
Sunower stalks
24% KOH1% NaHBH4
Wheat straw
24% KOH1% NaHBH4
Natural grass
Pigeon pea stalks
Green coconut
husks

4% KOH coupled with steam
21.8% of total dried raw

materials
materials
materials
Akpinar et al. (2009)

Samanta et al. (2012)
Samanta et al. (2013)
Jayapal et al. (2014)
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plants. Further to evaluate the utilization pattern of XOS by

rumen bacteria, Cotta (1993) attempted to produce XOS by
phosphoric acid. The effect of weak sulphuric acid (0.25 M) in
XOS production from corn cob xylan at 90 1C in different hours
of incubation has been reported recently (Samanta et al., 2012a).
Xylan is hydrolyzed to a number of oligosaccharides ranging
from X1 to X3 and above. The production of XOS is increased
along with increasing the duration of hydrolysis (Akpinar,
Erdogan, & Bostanci, 2009a). However, increasing the time of
hydrolysis is accompanied with higher degradation of xylan
into xylose and XOS. The reaction time for acid hydrolysis
beyond 30 min resulted in monosaccharide production, which
was in accordance with earlier ndings (Samanta et al., 2012a).
The time and concentration of acid played a very critical role in
the production of XOS.
5.2.
Enzymatic process
The conversion of xylan into value added useful products by

enzymatic process holds strong promise for the use of a variety
of un-utilized and under-utilized agricultural residues. Thus,
several attempts were made to produce XOS by enzymatic
means from xylan of diverse plant biomass origin including oat
spelt, beech wood, birch wood, corn cobs, sugarcane bagasse,
natural grass, oil palm frond etc. During adoption of enzymatic
means of XOS production, the enzyme preparations should
have lower exoxylanase activity so that xylose production
could be minimized in the XOS mixture. The yield of XOS
from xylan varies according to the source of xylan, enzyme
activity as well as incubation conditions such as pH, hydrolysis
time, temperature of incubation, etc. Exposure to xylanase
enzyme on xylan originated from corn cob, wheat bran, peanut
shell, oat spelt; the yield of XOS i.e. xylobiose (3.424.70 mg/ml),
xylotriose (0.343.66 mg/ml) and xylotetrose (nil to 1.99 mg/ml)
varied greatly (Yang, Yang, & Liu, 2007). While generating
prebiotic from alkali pre-treated sugarcane bagasse, Brienzo,
Carvalho, and Milagres (2010) noticed XOS yield in the range of
2.796.38 mg/ml while applying enzyme dose from 5 to 30 U/g
of substrate. The use of higher enzyme loading in the hydrolysis of wheat bran xylan resulted in the decrease of oligoxylosides yield along with reduced degree of polymerization.
Recently, Response Surface Methodology (RSM) analysis has
been applied for minimizing xylose and maximizing XOS from
natural grass, sugarcane bagasse (Samanta et al., 2012; Jayapal
et al., 2013). It indicated a maximum yield of xylobiose (11 g/
100 g xylan) could be possible at pH of 5.0, incubation temperature of 45.2 1C, enzyme dose of 17.4 U for a reaction time of
10.1 h. In order to obtain higher xylotriose (7.06 g/100 g xylan)
from the enzymatic hydrolysis of Sehima nervosum (natural
grass) xylan, the ideal conditions from response surface model
were pH 5.1, temperature 40.3 1C, enzyme dose 13.2 U and
reaction time 16.6 h.
5.3.
Autohydrolysis process
In the process of autohydrolysis, the xylan rich plant biomass is

subjected to processing with water in specialized equipment at
particular temperature and pressure (Parr Reactor). The xylan
(hemicelluloses) molecules are progressively broken down into
smaller molecules i.e. xylooligosaccharides under the inuence
of hydronium ions. The hydronium ions are derived from

autoionization of water that causes catalytic depolymerisation
of hemicelluloses to xylooligomers and xylose. Further cleavage
of acetyl groups to acetic acid also adds hydronium ion in the
mixture for further production of xylooligosaccharides (Garrote,
Dominguez & Parajo, 2002). XOS has been prepared by application of autohydrolysis from corn cobs (Samanta et al., 2012a),
wheat straw, rice husks, barley straw, almond shell (Nabarlatz,
Ebringerova, & Montane, 2007), oil palm frond (Sabiha-Hanim,
Noor, & Rosma, 2011). This process leaves cellulose and lignin
in the solid phase of reaction mixture. The maximum yield of
xylooligosaccharides by autohydrolysis could be achieved under
medium severity conditions (Moura et al., 2006). Increasing the
severity factor for processing leads to decreased degree of
polymerization and increasing decomposition of xylooligosaccharides into xylose. The disadvantage of this process is
requirement of specialized equipments and generation of
unwanted products in the XOS mixture.
6.
Biological role of xylooligosaccharides
Despite the description of prebiotic concept by Gibson and

Roberfroid (1995), the benecial effects of xylooligosaccharides
were realized by Imaizumi, Nakatsu, Sato, Sedarnawati, and
Sugano (1991), who noticed that supplementation of XOS at
the expense of starch or sucrose improved the diabetic
symptoms including reduction of liver triglycerides and phosphatidylcholine and recommended use of XOS as a substitute
for sugar in diabetic patient. The prebiotic efcacy of XOS
depends on its molecular weight (Hughes et al., 2007). There
are several benecial effects of prebiotics on host health.
These positive effects of XOS are linked with the optimization
of colonic functions as well as metabolism such as increase or
change in the composition of short chain fatty acids, increased
faecal weight and mineral absorption, immune stimulation,
decreased colonic pH, nitrogenous end products, procarcinogenic enzymes (Aachary & Prapulla, 2011; Samanta,
Jayapal, Senani, Kolte, & Sridhar, 2013a). Apart from prebiotic
activity, these bioactive molecules are believed to alleviate
disease symptoms in diabetes, cancer, stress (Ando et al. 2004;
Gobinath, Madhu, Prashant, Srinivasan, & Prapulla, 2010;
Wang, Cai, Wang, & Sun, 2011).
6.1.
Selective growth stimulation of gut microora
Based on the occurrence of microbial habitat in the gastrointestinal tract, all the vertebrates may be classied into two
categories viz.; Mono gut fermentor all non-ruminants (namely
human, poultry, swine, horse, rabbit, donkey etc.) and twine gut
fermentor all ruminants and pseudo-ruminants (namely
cattle, buffalo, sheep, goat, mithun, camel etc.). The gastrointestinal tract of ruminants is featured with the provision of two
microbial habitats i.e. foregut (rumen, reticulum and omasum)
and hindgut (caecum). On the other hand, non-ruminants are
featured with the presence of single microbial habitats i.e.
hindgut only (Samanta et al., 2013a). By virtue of unique
environment available in terms of nutrients, pH, interaction,
temperature, crosstalk inside the gastrointestinal tract, both
benecial and harmful microora gets their niches for growth
and multiplication (Samanta, Jayapal, Senani, Kolte, & Sridhar,

2011). Ignoring the substrate utilization pattern or biochemical
features, gut microora can be broadly grouped into two main
categories namely pathogenic or harmful and health promoting
or benecial. The class of pathogenic bacteria includes Escherichia coli, Streptococcus faecalis, Salmonella enterica, Clostridium
perfringens etc. Lactobacillus and Bidobacterium inhabiting within
the constituency of the gastrointestinal tract are presumed to be
the major members of benecial class. The signicance of
prebiotic consumption arises from their ability to selectively
stimulate the growth and multiplication of indigenous bidobacteria and lactobacilli in the hindgut which in turn suppresses
the activity of putrefactive or harmful bacteria and reduces the
concentration of toxic fermentation products in the gastrointestinal tract (Tomomatsu, 1994; Samanta et al., 2007, 2010).
XOS either in crude or puried form (generated from
Eleusine coracana bran) possesses prebiotic potentiality in
terms of selective stimulation of benecial gut microora.
At in vitro culture conditions, XOS stimulated the growth of
several probiotic strains such as Bidobacterium adolescentis
NDRI 236, Bidobacterium bidum NCDC 2715, B. bidum ATCC
29521, Lactobacillus brevis, Lactobacillus plantarum NDRI strain
184 after 48 h of incubation (Manisseri & Gudipati, 2012).
There is also concomitant increase in the dry cell biomass
of probiotic strains after 48 h of incubation. Among crude and
puried XOS, the latter is more potent and more effective to
exhibit growth stimulation of probiotic strains. Recently,
prebiotic potentiality of corn cob generated XOS was assessed
by enumerating the colony forming units for four proven
probiotic strains. It revealed that prebiotic action of XOS was
higher with probiotics Enterococcus faecium followed by Enterococcus fecalis, Lactobacillus maltromicus, Lactobacillus viridiscens;
implicating variable growth stimulatory effect of XOS
(Samanta et al., 2012). In case of arabinoxylan fractionated
from wheat bran subjected to fermentation by human fecal
microora, there is increased proliferation of bidobacteria,
lactobacilli and eubacteria (Hughes et al., 2007). The molecular weight of arabino-xylan has inverse relation with prebiotic
effects. The prebiotic ability of arabino-xylan, particularly
selective growth stimulation of Bidobacterium and Lactobacillus, increases with the decreased degree of polymerization.
In line with the in vitro fermentation prole of XOS, these
bioactive molecules are found to exhibit selective growth
stimulation of benecial gut microora even in laboratory
animals. Oral administration of XOS to rat and mice signicantly increased the moisture content of faeces, total caecum
weight and population of bidobacteria in addition to reduced
caecum pH (Chung, Hsieh, & Chan, 2002; Chan, Chung, Chang,
& Chan, 2005). Both healthy and diabetic rats maintained on
diet containing either 5 or 10% XOS exhibited signicant
increase in caecal bidobacteria and lactobacilli population
(Gobinath et al., 2010). The increased population of benecial
gut microora is accompanied with several changes in the gut
such as decrease in caecum pH and increase in total weight of
caecum. The reduction in the pH of caecal content might be
attributed to the increase in the concentration of short chain
fatty acids generated from the selective fermentation of prebiotics by the benecial gut microora such as bidobacteria
and lactobacilli (Gibson & Roberfroid, 1995). With regard to
selective growth stimulating potentiality between XOS and FOS,
67
the former has been found to be more bidogenic, while the

latter is considered to be more specic to lactobacilli (Okazaki,
Fujikawa, & Matsumoto, 1990; Klessen, Hartmann, & Blaut,
2001; Gobinath et al., 2010; Chapla, Pandit, & Shah, 2012).
Evidences suggest that prebiotic XOS also possess anticarcinogenic properties. Application of XOS signicantly
reduces the precancerous lesions induced by 1,2-dimethylhydrazine in rats and supplementation of XOS was more effective
than fructooligosaccharides in stimulating the growth of bidobacteria (Hsu, Liao, Chung, Hsieh, & Chan, 2004). Emergence
of numerous applicability reports of XOS throughout the world
indicates the growing consumer awareness for such important
biomolecules of lignocellulosic biomass origin. Albeit, maximum levels of XOS depend on various factors, the recommended dose of XOS is 0.12 g/kg body weight in adults (Oku &
Sadako, 2002). Consumption of XOS also found to restrain the
growth and multiplication of pathogenic and putrefactive gut
microora (Mussatto & Manchilha, 2007; Samanta et al., 2013).
Prebiotics XOS are also able to ensure relief from gastrointestinal disorders such as constipation, inammatory bowel disease, gastritis etc. through restoration of gut microora
homeostasis (Rautonen, Apajalahti, Kettunen, & Sikanen,
2004; Gibson & Beer, 2004; Carvalho, Neto, Silva, & Pastore,
2013). Chung, Hsu, Ko, and Chan (2007) studied the daily dosing
of XOS in elderly with average age above 75 years belonging to
both male and female sex. The human studies revealed
signicant decrease of fecal pH with concomitant increase of
fecal moisture and population of bidobacteria. Further, the
studies did not nd any inuence of XOS on the different
gastrointestinal behaviour or symptoms such as appetite,
rumbling, voices of gut, belching, atulence, faecal smelling
etc. In addition to above studies, consumption of XOS by
pregnant woman is recorded to be highly effective against
constipation without any other adverse effects (Tateyama
et al., 2005).
6.2.
Antioxidant property of XOS
Owing to the radical changes of lifestyle, stress has become a

great concern to the present human society. Modern management approaches (such as weaning) in livestock also cause
stress of different degree which adversely affect the subsequent productivity of animals. In the event of stress, there is
breakdown in the equilibrium status between reactive oxygen
species (ROS) and antioxidant systems. This results into
accumulation of excess ROS within the body and causing
damage to various biomolecules such as DNA, protein,
carbohydrates, membrane lipids leading to cell death and
tissue injuries (Aruoma, 1998). As a result, there is manifestation of variety of chronic health problems such as cardiovascular disease, arthritis, diabetes, cancer, Alzheimers disease,
neurological disorders in addition to the acceleration of aging
process. Recently, there are few reports on the antioxidant
activity of oligosaccharides such as soya-oligosaccharides,
XOS, etc, which could effectively be used to ameliorate stress.
Rats with streptozotocin (STZ) induced diabetes, exhibited
increased plasma concentration of antioxidant enzymes such
as catalase and glutathione reductase while receiving XOS at
10% dietary concentration (Gobinath et al., 2010). The antioxidant activities of XOS are accompanied by reduced plasma
68
glucose, cholesterol and creatinine levels in experimentally

induced diabetes in rats. Consumption of high fat diet results
into hyperlipidemia coupled with increased oxidative stress
(Sreekumar et al., 2002). Diet containing 5% XOS is recorded to
alleviate the symptoms of oxidative stress resulted from high
fat diets. The supplementation of XOS in rats maintained on
high fat diet, reduced oxidized glutathione activity and
increased the contents of reduced glutathione, superoxide
dismutase, catalase, glutathione peroxidise in serum, liver
and heart (Wang et al., 2011). The precise mechanism of
action of XOS in reduction of adverse affects of oxidative
stress is yet to be substantiated through scientic ndings.
The source of XOS also inuences the antioxidant potentiality of the particular biomolecule. Antioxidant assay techniques (DPPH and FRAP assay) have shown that XOS from ragi
bran at 10 mg exhibited antioxidant activity of 12% and
reached up to 70% at 60 mg concentration; indicating higher
antioxidant potential of XOS generated from rice, maize or
wheat (Veenashri & Muralikrishna, 2011).
6.3.
Short chain fatty acid production
Due to the presence of -1,4-xylosidic bonds, XOS is not

digested by the enzymes secreted by vertebrates own organelles and reaches at hindgut after retaining its structural
integrity. Hindgut is the site of microbial habitat in body
containing both benecial and harmful microora. The XOS is
selectively utilized by benecial microora especially the bidobacteria for growth and multiplication resulting into the
production of short chain fatty acids viz.; acetic, propionic and
butyric acids (Macfarlane & Macfarlane, 2003; Manisseri &
Gudipati, 2012). The efcient and complete utilization of XOS
requires the concerted activities of several enzymes such as
-xylosidase, -glucoronosidase, -L-arabinosidase, acetyl esterase. Under culture conditions, Bidobacterium adolescentis, Bidobacterium infantis and B. bidum are found to secrete only
-xylosidase and -arabinosidase for growth and multiplication
(Zeng, Xue, Peng, & Shao, 2007). There is wide variation on the
utilization of XOS among the different strains of rumen bacteria
such as Selenomonas ruminantium and acetic, lactic and propionic
acids are the major end products of fermentation (Cotta &
Whitehead, 1998). The degradability of XOS varies at different
segment of hindgut in pigs. The microbial inoculums of caecum
and colon ferments XOS at faster rate than the inoculums of
ileum as it is evidenced by higher accumulation of short chain
fatty acids (Moura et al., 2008). Lactate is an intermediary
product of XOS at the beginning of fermentation and in due
course of time this is further metabolized into acetate, propionate and butyrate by common intestinal microora. Switching
the growth stimulation of benecial gut microora by consumption of prebiotics is associated with decrease in fecal and
caecal pH, followed by reduction in the population of harmful
microora such as C. perfringens, E. coli etc. The concentration of
putrefactive substances is also reduced in the stool and urine as
a result of consumption prebiotics (Bornet, Brouns, Tashiro, &
Ouvillier, 2002). The bidogenic activity is associated with the
reduction in concentration of fecal enzymes such as -glucuronosidase, nitroreductase, azoreductase, glyocholic acid hydrolase responsible for generation of harmful metabolites at the
hindgut (Samanta et al., 2013). Curbing the secretion of these
undesirable enzymes has additional benets in terms of dropping the production of noxious substances such as vasoconstricting amines, phenols, steroid metabolites, bacterial
toxins, carcinogens etc at the hindgut.
6.4.
Immune stimulation
There is growing interest for enhancing immune status

through administration of prebiotics especially during disease conditions or in vulnerable subjects such as in early age
groups or elderly people. Albeit, frugal information is available on how the host gut microora recognize ingested
prebiotics for exhibiting immune-modulating properties, yet
these bioactive substances or their metabolites are found to
be closely associated with the gut associated lymphoid
tissues (GALT); major contributor of intestinal immune functions (Saad, Delattre, Urdaci, Schmitter, & Bressollier, 2013).
Alternatively, the innate defence responses are activated
following interaction between sugar molecule and innate
receptors present at macrophages and dendritic cells
(Arnold, Dwek, Rudd, & Sim, 2006; Nezlin & Ghetie, 2004).
Evidently oral administration of prebiotics have elucidated to
positively modulate diverse parameters of immune systems
such as secretion of IL-10, interferon, NK cell activity, lymphocyte proliferation, polymeric immunoglobulin receptors
expression in both intestine and colon of mice (Hosono et al.,
2003; Hoentjen et al., 2005; Nakamura et al., 2004).
6.5.
Lipidaemic effects
Studies in laboratory animals evidenced the potential capability

of prebiotic XOS on reduction of serum triglycerides and
cholesterol levels; important markers for assessing vulnerability
of cardiovascular diseases. Before the emergence of the concept
of prebiotics, Imaizumi et al. (1991) noticed signicant reduction
of serum cholesterol and triglycerides in diabetic rats following
administration of XOS. High fat diet results into abnormal blood
lipid prole; very common in present century. Incorporation of
5% XOS in rats (maintained on high fat diets) signicantly
improves the lipid levels indicated by signicant drop in
triglycerides, total cholesterol, low density lipoprotein and
increase in high density lipoprotein levels (Wang et al., 2011).
XOS induced lipidaemic responses are also noticed in type 2
diabetes mellitus subjects (Sheu, Lee, Chen, & Chan, 2008).
Recently, similar lipidaemic effects were also reported in broiler
birds (Samanta et al., 2013b). However, potentiality of XOS as a
modulator of lipid metabolism on human subjects is still
contentious issue. A slight but signicant reduction of serum
cholesterol and triglycerides was noticed among young woman
after regularly dosing with 2.7 g of XOS (Na & Kim, 2007); while
consumption of 3.8 g XOS per day failed to evoke lipidaemic
effects on elderly subjects (Chung et al., 2007). Although the
mechanism of lipidaemic response of XOS is still not understood, some school of thought believes in the role of propionate
generated from the fermentation of XOS by gut benecial
microora. Propionate inhibits lipogenesis in liver (Wolever,
Spadafora, Cunnane, & Pencharz, 1995; Demigne et al., 1995).
Alternately, the colonic propionate or rest short chain fatty
acids stimulate production of intestinal glucagon like peptide 1
(GLP-1) to promote insulin synthesis and thereby cause
inhibition of lipolysis and stimulation of glycogen synthesis

(Frost, Brynes, Dhillo, Bloom, & McBurney, 2003; Meier et al.,
2006; Delzenne, Cani, & Neyrinck, 2007).
7.
Safety aspects of XOS
Of late, many patents pertaining to the process of XOS

production utilizing a range of raw materials by chemical,
enzymatic or autohydrolysis processes have evolved across
the world. Owing to its multidimensional activities, XOS
could be incorporated in many food ingredients in order to
ensure proper gut health and functionality. Further, ban
(within European Union) or restriction (in India) on feed
application of antibiotics had raised the hope for greater
application of prebiotics to guard the gastrointestinal tract
from unwanted effects of pathogenic microora. Nowadays,
nobody raises the safety concern for application of inulin or
galacto-oligosaccharides in a wide range of products including chocolate, biscuits, bread, etc. due to their history of safe
usage and inherent ability to address the human health
concerns. In animal models, administration of XOS in the
range of 710% of diets does not exhibit any adverse effects
(Hsu et al., 2004; Santos, San Mauro, & Diaz, 2006; Gobinath
et al., 2010). However, studies on human subject revealed
benets of XOS provided the daily consumption is between 2
and 5 g (Kobayashi, Okazaki, Fujikawa, & Koga, 1991). The
maximum permissible dose that does not lead to any gastrointestinal symptoms is 0.12 g/kg body weight in humans (Oku
& Sadako, 2002). Preliminary investigations elucidates the
great potentiality of XOS in terms of bidogenic activity vis a
vis its effect on other associated functions such as immune
stimulation, enhanced mineral absorption, cholesterol, glucose reducing ability, etc. This opens the door for wider
applicability of XOS in diversied products.
8.
Summary
Amongst the nutraceuticals, the indigestible compounds (prebiotics) having potential to selectively stimulate the growth and
multiplication of benecial gut microora is receiving greater
attention around the world. The acceptability of prebiotics by
consumers is due to their additional technological features
including lower caloric values, non-carcinogenic, fat replacing
ability, stability under wider pH range, acceptable taste, etc. XOS
occupies signicant niche amongst the prebiotics as it addresses the issues such as effective disposal of agricultural
wastes, value addition, taking care of human and livestock
gut health, fortication of food items, alternative feed additives
in livestock and so on. Presently, the prebiotic production is
largely dependent on the crops mostly used for human consumption (Jerusalem artichoke, chicory root, asparagus, etc.)
resulting in higher cost of production for the nal product i.e.
prebiotics. Under such circumstances, approach to explore
agricultural crop residues as initiator for nutraceuticals production might pave way for the manufacturers to substantially
reduce the cost of raw materials as well as solve the disposal
problems. Further, realizing the close link between food and
health; present day consumers are much more inclined
69
towards biomolecules of natural origin that have attributes like

disease prevention and health promotion in addition to nutritional importance. The research and development for novel
process or raw materials towards XOS production is continuing.
Future stance of XOS lies in its economic production on an
industrial scale and their validation through animal models or
human clinical trials. Furthermore, studies need to be carried
out to analyze the benecial effects of XOS fortication/supplementation through application of advanced biological tools.
Acknowledgement
The authors are thankful to the Department of Biotechnology,
Ministry of Science and Technology, Government of India for
providing the nancial assistance (Grant no. BT/PR5703/AAQ/
1/504/2012) to carry out the present work.
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Bioactive Carbohydrates and Dietary Fibre 5 (2015) 62 71

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