prebiotics and probiotics science and technology || prebiotic potential of xylo-oligosaccharides

8 Prebiotic Potential ofXylo-OligosaccharidesH. Makelainen . M. Juntunen . O. Hasselwander

8.1 Introduction

Xylo-oligosaccharides (XOS) are chains of xylose molecules linked with b1–4bonds (> Figure 8.1) with degree of polymerization ranging from 2 to 10.

XOS are naturally present in fruits, vegetables, bamboo, honey and milk, and

can be produced at industrial scale by enzymatic hydrolysis from xylan, which is

the major component of plant hemicelluloses and therefore readily available in

nature (Alonso et al., 2003; Vazquez et al., 2000).

XOS are non-digestible carbohydrates and have been suggested to exert

prebiotic activity. They were hence first used as food ingredient for gastrointesti-

nal health in the 1990s in Japan. This chapter provides an overview of XOS with a

particular focus on the prebiotic potential.

8.2 Manufacture of XOS

XOS can be produced commercially by hydrolysis of xylan, the most abundant

hemicellulosic polymer. Possible lignocellulosic raw materials for XOS produc-

tion include corn cobs, hardwoods, straws, bagasses, hulls, malt cakes and bran.

Xylan can either be hydrolyzed enzymatically, by chemical methods (hydrother-

mal treatments) or a combination of both. The resulting crude XOS solutions

require a sequence of purification steps to yield high purity XOS containing at

least 70–95% XOS (Moure et al., 2006; Vazquez et al., 2000). Depending on the

source of raw material, XOS may be branched and contain arabinose units or

carry acetyl or uronic acid residues (Vazquez et al., 2000).

XOS have been used as food ingredient predominantly in Asia, particularly

Japan and Suntory Limited (Japan) was the first commercial-scale producer

# Springer ScienceþBusiness Media, LLC 2009

applying enzymatic hydrolysis of xylan. The production volume of XOS was

estimated at 650 tons annually in 2004 and the XOS price was 2,500 Yen/kg

(Taniguchi, 2004). More recently, XOS were also offered by the Chinese pro-

ducer Shandong Longlive Bio-technology co. In 2003, XOS represented only a

small proportion (less than 3%) of the total Asian oligosaccharide market

(Nakakuki, 2003).

The commercial XOS products are available in syrup or powder form and are

predominantly composed of the disaccharide xylobiose and the trisaccharide

xylotriose with small amounts of higher oligosaccharides also present.

8.3 XOS as Prebiotics

8.3.1 Resistance to Digestion

XOS are relatively stable in acidic conditions due to structural properties.

This may endow protection from decomposition when passing through the

stomach (Imaizumi et al., 1991). The degradation of XOS (xylobiose) in the gas-

trointestinal tract has been studied in vitro with an artificial model of digestive

enzymes (a-amylase, pancreatin, gastric juice and intestinal brush border

enzymes) and no hydrolysis of xylobiose was observed (Koga and Fujikawa,

1993; Okazaki et al., 1991).

The fate of xylobiose was also studied in humans after oral administra-

tion. Xylobiose was not excreted into feces or urine during 24-h following

the ingestion, thus, supporting the fact that xylobiose is degraded in vivo

not by the action of digestive enzymes, but by the gastrointestinal microbiota

(Okazaki et al., 1991).

. Figure 8.1Disaccharide Xylobiose (b-D-xylopyranosyl (1!4)- D-xylopyranose).

246 8 Prebiotic Potential of Xylo-Oligosaccharides

8.3.2 Fermentation by the GastrointestinalMicrobiota and Selective Stimulation of Growthand/or Activity of Intestinal Bacteria Associatedwith Health and Well-Being

8.3.2.1 Pure Culture Studies

Pure culture fermentation studies with single microbes and substrates can be used

to identify bacterial strains that are able to degrade oligosaccharides. Since these

fermentations do not resemble the competitive environment of the colon, the

results can be used to gain knowledge of the fermentative capacity of individual

strains within the intestinal microbial population, but not to study the effects that

oligosaccharides have on the whole microbiota.

XOS are reported to be preferentially fermented by a relatively limited

number of intestinal microbes in vitro. Several pure culture studies have indicated

that XOS are well utilized by Bifidobacterium species, namely some strains of

B. bifidum, B. catenulatum, B. longum, B. animalis and B. adolescentis (Crittenden

et al., 2002; Jaskari et al., 1998; Moura et al., 2007; Palframan et al., 2003; Yamada

et al., 1993). Furthermore, utilization of XOS seems to be strain-dependent, since

not all studies have shown enhancement of for example all B. longum and

B. adolescentis strains (Hopkins et al., 1998). Lactobacilli are not able to utilize

XOS as a sole carbon source (Jaskari et al., 1998; Kontula et al., 1998), with the

exception of Lactobacillus brevis, which growth was enhanced moderately by XOS

(Crittenden et al., 2002; Moura et al., 2007). Some other intestinal microbes are

also able to utilize XOS, but not to the same extent as bifidobacteria. Crittenden

and co-workers studied the fermentation of a wide group of numerically domi-

nant saccharolytic intestinal bacterial species and demonstrated that besides

many Bifidobacterium strains only some Bacteroides isolates were efficiently

fermenting XOS. Escherichia coli, enterococci, Clostridium difficile and Clostridi-

um perfringens were not able to ferment XOS (Crittenden et al., 2002). Jaskari

et al. found that XOS was metabolized by bifidobacteria, but also moderately by

Bacteroides thetaiotaomicron, Bacteroides vulgatus and Clostridium difficile. How-

ever, these strains mainly utilized the monosaccharide xylose fraction of the XOS

mixture, which in humans does not reach the colon (Jaskari et al., 1998).

Unpublished research (carried out by M. Juntunen at Danisco’s Health and

Nutrition Center in Kantvik, Finland) suggests a selective utilization of XOS by

Bifidobacterium lactis strains (> Figure 8.2). Other tested microbes showed poor

growth on commercial (XOS Longlive 95P, Shandong Longlive Bio-technology

Prebiotic Potential of Xylo-Oligosaccharides 8 247

.Figure

8.2

Thegrowth

ofaselectionofbifidobacteria,lactobacilliandothermicrobialstrainsonXOS.T

hegrowth

wasmeasuredasthech

angein

absorbance

(600nm)ofliquid

samplesandrepresentedtheareaunderthegrowth

curve(O

Dxminutes)obtainedduringthe24-h

growth

exp

eriment(Jaskarietal.,1

998).Theareaoftheco

ntrolmedium

withoutaddedcarbohydrateswassubtractedfrom

results.


co., China) and hardwood-derived XOS (XOS dp2 and XOS dp2–10) when

compared to growth obtained on glucose during 24-h incubation.

Interestingly, pure culture studies have also shown that for a number of

Bifidobacterium strains the bacterial growth was higher on oligosaccharides in

comparison to their monosaccharide constituents (Hopkins et al., 1998), thus,

indicating that there may be a specific membrane transport mechanism for XOS

but not xylose. Some bifidobacteria might import XOS before hydrolyzing it

(Crittenden et al., 2002), which could offer a competitive advantage against cross-

feeding by other microbes in the intestine.

8.3.2.2 Batch Culture Studies

Effects of oligosaccharide fermentation on predominant intestinal microbial groups

can bemonitored in batch culture fermentations and continuous or semi-continuous

color simulations using colonicmicrobiota from fecal samples. The effects of different

prebiotic oligosaccharides (FOS, inulin, lactulose, galacto-oligosaccharides (GOS),

isomalto-oligosaccharides (IMO), soybean oligosaccharides (SOS), and XOS) on

microbiota were compared in batch fermentations (Rycroft et al., 2001). Fermenta-

tion properties of different oligosaccharides varied and resulted in different

responses in the composition and activity of microbiota. Fermentation of all

oligosaccharide compounds increased the numbers of bifidobacteria and most

decreased clostridia, but XOS and lactulose produced the highest increase in

bifidobacteria, whereas FOS were the most effective substrate for lactobacilli. A

similar significant increase in numbers of bifidobacteria, and also lactobacilli, was

seen as a result of XOS fermentation in a semi-continuous simulator system

inoculated with adult feces (Zampa et al., 2004). Microbial fermentation of XOS

moderately increased the production of gases, and the concentrations of lactate,

acetate, and propionate in both studies (Rycroft et al., 2001; Zampa et al., 2004).

Zampa and co-workers also reported beneficial effects of XOS not only derived

from increased populations of bifidobacteria and lactobacilli, but also from

reduced concentrations of secondary bile acids, which exert negative actions on

the colon and present a dose dependent toxic potential related to their co-muta-

genic and tumor-promoting properties (Moure et al., 2006).

8.3.2.3 Animal Studies

The bifidogenic effects of XOS have also been observed in animal studies

(> Table 8.1). Studies in rats have demonstrated that XOS significantly stimulate


the growth of cecal and fecal bifidobacteria. Furthermore, XOS induced an even

larger increase in bifidobacteria numbers than an equivalent dose of FOS, which

had a greater effect on the Lactobacillus population (Campbell et al., 1997; Hsu

et al., 2004). A more recent feeding-trial with mice (Santos et al., 2006) compared

the long-term effects of various prebiotics on the microbial populations. In

this study, 1% of FOS, inulin, lactulose, XOS, SOS, IMO, or transgalacto-

oligosaccharides (TOS) were administered to the basal diet and the effects on

small and large intestinal microbiota were determined after 6 months of inter-

vention. From all the prebiotics tested, XOS was the most efficient substrate to

. Table 8.1

Animal studies on effects of XOS

Animal studies Major findings Reference

Diabetic rats (n = 8), dietary sucroseand corn starch replaced with XOS for 5weeks

Improvement in diabetic symptomssuch as elevated serum glucose,cholesterol and triglycerides

Imaizumiet al.(1991)

Rats (n = 50) were fed with fiber-freediet, or diet containing 7.5% of oatfiber, gum arabic, FOS or XOS for 17days

Oligosaccharides and gum arabic allincreased the cecal wall and contentsweight and decreased cecal pH withconcomitant increase in SCFA.Excretion of nitrogen in feces wasincreased, thus, blood urea and renalnitrogen excretion decreased

Youneset al.(1995)

Rats (n = 44) and mice (n = 52) wereassigned to control, FOS, XOS or gumarabic group for 14 days

XOS did not affect microbiota ofanimals, and only moderate effects oncecal cell proliferation were found. FOSincreased significantly bifidobacteria inrats

Howardet al.(1995)

Rats (n = 10 in each group) were fedwith control, cellulose, oligofructose,FOS or XOS containing diet for 14 days

Fecal pH was lowest and bifidobacterianumbers highest in the XOS group.Consumption of XOS increased mostthe weight of colon and cecum

Campbellet al.(1997)

Rats (n = 40) were fed with basal, XOSor FOS diet for 35 days, and treatedwith 1,2-dimethylhydrazine (DMH) toinduce colon carcinogenesis

Oligosaccharides decreased cecal pH,increased cecal weight andbifidobacteria population, but XOS hada stronger effect than FOS. Botholigosaccharides reduced theformation of precancerous lesion

Hsu et al.(2004)

Mice (n = 16 in each group), dietsupplemented (1%) with nine differentoligosaccharides for a 6- month studyperiod

XOS was most efficient prebiotic inincreasing lactobacilli andbifidobacteria counts, and in reducingsulphite-reducing clostridia

Santoset al.(2007)


increase bifidobacteria, lactobacilli and total anaerobic microbial numbers in the

colon. Of all the oligosaccharides, XOS supplementation also resulted in

the strongest reduction of sulphite-reducing clostridial strains, thus affecting

the colonic microbiota in an overall favorable manner.

8.3.2.4 Human Intervention Studies

Reports from few human interventions, mostly conducted in Japan, regarding

colonic fermentation of XOS have been published to date (> Table 8.2). Okazaki

and co-workers fed 5 g of XOS daily for 3 weeks to healthy men, and found

a significant increase in fecal bifidobacteria numbers. The effect of XOS

. Table 8.2

Human intervention studies conducted on XOS

Human intervention study Major findings Reference

Healthy men (n = 9), 5 g/day XOS for3 weeks

Significant increase in Bifidobacteriumand Megasphaera, other microbesunaffected. Increased fecal acetic acidand decreased pH

Okazakiet al.(1990a)

Healthy men (n = 5), 1 and 2 g/dayXOS for 3 weeks each

Significant increase in Bifidobacterium,other microbes unaffected, Consumptionof 2 g was more effective, but even doseof 1 g of XOS increased bifidobacteria

Okazakiet al.(1990b)

Healthy men (n = 10), 2, 5 and 10 g/day XOS

Occurrence of diarrhea decreased withconcomitant increase in Bifidobacteriumwith daily doses of 2 and 5 g

Kobayshiet al.(1991)

Healthy men (n = 9), 5 g/day XOS for3 weeks

Putrefaction products in feces (p-cresol,indole, skatol) decreased

Fujikawaet al.(1991)

Constipated women (n = 40),0. 4 g/day XOS for 4 weeks

Defecation frequency and stool quantityincreased, self reported quality of lifeimproved

Iino et al.(1997)

Constipated pregnant women(n = 29), 4.2 g/day XOS for 4 weeks

XOS effective in reducing severeconstipation and normalizing stoolconsistency. Clinical symptoms scoresimproved

Tateyamaet al.(2005)

Healthy elderly (n = 9 in control group;n = 13 in XOS group), 4 g/day XOS for3 weeks

Significant increase in Bifidobacteria andfecal moisture content, decreased fecalpH

Chunget al.(2007)


administration on the growth of several other microorganisms was also studied,

and only Megasphera numbers were significantly affected (increased) in addition

to bifidobacteria, indicating a very selective proliferation effect (Okazaki et al.,

1990a). Fecal pH reduced and acetic acid concentrations increased during this

intervention. In another trial by the same researchers, Bifidobacterium numbers

were increased in the feces of subjects consuming only 1 and 2 g of XOS per day,

although the higher daily dose resulted in a more significant increase (Okazaki

et al., 1990b). Even as low a daily dose as 0.4 g of XOS was shown to increase the

numbers of bifidobacteria in human fecal samples, without effects on other

microbial groups (Iino et al., 1997). Furthermore, decreased concentrations of

putrefactive products such as p-cresol, indole and skatole were measured con-

comitantly with an increased ratio of fecal bifidobacteria (Fujikava et al., 1991). In a

more recent human intervention study in elderly subjects (Chung et al., 2007), a

daily XOS dose of 4 g increased significantly the Bifidobacterium numbers from

106 to 108 cfu/g of wet feces during a 3 week study period. Clostridium perfringens

levels remained unchanged and the fecal pH was decreased and fecal moisture

content increased during the study period. No difference in adverse gastrointes-

tinal symptoms (flatulence, discomfort, stool consistency, defecation frequency)

between the control and study group was recorded, suggesting that XOS was well

tolerated by the elderly.

The limitations of the human interventions studies carried out to date are

relatively small number of subjects, the (in most cases) uncontrolled study-design

and the use of plating method in enumeration of microbes in feces, thus, larger

controlled human intervention studies are needed to confirm the prebiotic status

of XOS.

8.3.2.5 Effect of Substitution and Origin of XOS

The effect of differently substituted XOS on fermentability was first recognized by

Van Laere et al., who included a linear XOS and arabinoxylo-oligosaccharides

(AXOS) in a fermentation study of complex plant cell wall derived oligosacchar-

ides (Van Laere et al., 2000). It was shown that the linear XOS were fermented by

more of the tested intestinal microbial strains tested compared to the branched

AXOS. The number and nature of the substitutes in XOSmolecule appears to affect

the fermentation speed and the metabolites produced. This was in particular

demonstrated in a batch fermentation study (Kabel et al., 2002), where non-

substituted XOS (nXOS) and arabino-XOS (AXOS) were reported to be fermented


more quickly than structurally more complex acetylated XOS (AcXOS) and XOS

containing a 4-O-methylglucuronic acid group (GlcA(me)XOS). In this study, the

fermentation of the least substituted molecules XOS and AXOS resulted in quick

production of acetate and lactate, whereas the fermentation of AcXOS and GlcA

(me)XOS increased the production propionate and butyrate. These results highlight

the importance of detailed elucidation of the structural features of non-digestible

oligosaccharides in relation to their fermentation properties.

8.3.3 Effects on Health

The formation of preneoplastic lesions (aberrant crypt foci, ACF) in the distal

colon is used as a biomarker for colon carcinogenesis. In a study with rats (Hsu

et al., 2004), the ACF formation was induced with 1,2-dimethylhydrazine (DMH,

a tumor promoter) and the rats were fed with basal diet or diet containing XOS or

FOS. Dietary supplementation with both oligosaccharides inhibited the develop-

ment of precancerous colonic lesions and simultaneously lowered the cecal pH

level through increased concentrations of short chain fatty acids (SCFA) and

increased the Bifidobacterium population and cecal weight. The decreased ACF

and increased cecal weight could be due to the normalization of epithelial cell

proliferation via increased concentrations of SCFA from oligosaccharide fermen-

tation by Bifidobacterium. In this study, XOS was more efficient in increasing the

bifidobacteria numbers and relative colonic and cecal wall weight than FOS,

leading the investigators to conclude that XOS could be more effective in increas-

ing and controlling the epithelial cell proliferation through a trophic effect of

produced SCFA. These findings are supported by other animal trials (Campbell

et al., 1997; Howard et al., 1995; Younes et al., 1995), although Howard and co-

workers found only moderate effects of XOS on cell density and cecal crypt

depths in mice, and no effects on microbiota (Howard et al., 1995). Campbell

et al. showed increased cecal and colonic weight in rats together with decreased

pH, increased SCFA (especially lactate and acetate) and bifidobacteria as a result of

XOS and FOS supplementation. In the same study, XOS supplementation resulted

in more significant changes in measured parameters than FOS supplementation.

Younes and co-workers found an increase in total cecal weight (wall and content

weight) after rats consumed XOS and FOS. The two oligosaccharides reduced the

pH in cecum significantly more than oat fiber or control diet, and increased cecal

SCFA concentrations. The ratios of SCFA differed considerably between oligosac-

charides with XOS producing more acetate and FOS butyrate. Also, fecal nitrogen


excretion increased and urinary nitrogen excretion decreased as a results of dietary

oligosaccharide supplementations (Younes et al., 1995). Results from these animal

trials indicate that fermentation of XOS could play a role in maintaining normal

mucosal differentiation and, thus, the integrity of the colonic mucosa.

A study on diabetic rats found that when simple carbohydrates in the diet

were replaced with XOS (10 g per day), the increased serum cholesterol and

triglyceride levels seen in diabetes were reduced and liver triglycerides levels to a

comparable level seen in healthy rats (Imaizumi et al., 1991). The researchers

concluded that XOS could be applicable to foods as a sweetener replacing sucrose,

which would benefit diabetic patients. However, these findings reported in animal

trials have not been confirmed in humans yet.

In some human interventions, effects of XOS on intestinal microbiota and

gastrointestinal function have been studied at the same time. Intervention in

constipated pregnant women showed marked improvements in the defecation

frequency and stool consistency during and after dietary supplementation with

4.2 g of XOS for 4 weeks. Before XOS administration the subjects recorded, in

average, to defecate only once per week. During and after the intervention,

defection frequency increased significantly to 6–7 times per week with concomi-

tant improvement in subject’s self-reported symptoms (Tateyama et al., 2005).

Similar findings with adult women have been reported previously (Iino et al.,

1997). Defecation frequency and abdominal symptoms improved simultaneously

with increased bifidobacteria numbers, and persisted 2–4 weeks after XOS inges-

tion was completed. Kobayashi et al. found that administration of 2 g of XOS per

day decreased the frequency of diarrhea in men (Kobayashi et al., 1991).

8.4 Safety and Regulatory Status

XOSwere tested for mutagenicity, acute and subchronic toxicity. XOS were found

to be non-mutagenic and showed no acute toxicity. Safety was also confirmed in a

90-day subchronic toxicity study in rats. These studies are mentioned in a

Suntory product brochure and were conducted with Suntory’s Xylo-oligo70

product (Biotec Suntory – Xylo-oligosaccharide brochure).

Limited data on digestive tolerability are available; however, volume of gas

produced by human fecal bacteria during 24-h fermentation in vitro was similar

for FOS and XOS (Rycroft et al., 2001) and no adverse effects have been reported in

the human interventions carried out in healthy subjects (Okazaki et al., 1990a,b)

including pregnant women (Tateyama et al., 2005) and elderly (Chung et al., 2007).


In Japan, XOS are approved as ingredients for Foods for Specified Health

Uses (FOSHU), specifically for foods to modify gastrointestinal conditions at a

recommended daily dose of 1–3 g (Japanese Ministry of Health, Labour and

Welfare, www.mhlw.go.jp/english/topics/foodsafety/fhc/02.html). Use of XOS as

food ingredient outside Japan and other Asian countries where XOS-containing

products are currently marketed may require specific regulatory approval.

8.5 Market Information and Application

XOS were first used as food ingredient in the 1990s in Japan and since 1997, 32

product launches (including new formulations and varieties) have been recorded

by Mintel’s global new products database (Mintel, 2008). Use of XOS in dietary

supplements accounted for 38% of all launches, use in dairy for 25%, use in sugar

and gum confectionery for 16%, use in non-alcoholic beverages for 13% and use

in baby food and soup for the remaining 8%. XOS is predominantly used in Asia

(Japan, China, South Korea, Vietnam, Taiwan) with Japan accounting for more

than half of the new product launches since 1997.

According to Suntory’s product brochure (Biotec Suntory – Xylo-

oligosaccharide brochure), XOS are acid- and heat-resistant. XOS remain intact

after heating for 1 h at 100�C within a pH range from 2.5 to 8. Heat stability has

also been confirmed at 120�C. Sweetness is claimed to be approximately 40% of

sugar after comparable refinement and viscosity should allow easy use in various

food applications (Biotec Suntory – Xylo-oligosaccharide brochure).

8.6 Summary

� XOS are resistant to digestion and are fermented by gastrointestinal microbiota.

� Fermentation of XOS by microbiota increases the concentrations of SCFA in colon,

especially acetate, propionate and lactic acid. Gas volumes produced are moderate.

� Results from in vitro and in vivo studies are promising and suggest that XOS may

selectively stimulate growth and/or activity of intestinal bacteria associated with

health and well-being, particularly bifidobacteria.

� Additional controlled human intervention studies using molecular techniques to

determine changes in fecal microbiota are needed to confirm the prebiotic potential

of XOS and the efficacious dose, which has been suggested to be as low as 1 g/day.

� XOS can be considered an emerging prebiotic, as the scientific evidence is still not

sufficient to classify XOS as an established prebiotic compound (Gibson et al., 2004).


http://www.mhlw.go.jp/english/topics/foodsafety/fhc/02.html

List of Abbreviations

ACF aberrant crypt foci

AcXOS acetylated xylo-oligosaccharides

AXOS arabinoxylo-oligosaccharides

DMH 1,2-dimethylhydrazine

dp degree of polymerization

FOS fructo-oligosaccharides

GlcA(me)XOS xylo-oligosaccharides containing a 4-O-methylglucuronic acid

group

GOS galacto-oligosaccharides

IMO isomalto-oligosaccharides

nXOS non-substituted xylo-oligosaccharides

OD optical density

SCFA short chain fatty acids

SOS soybean oligosaccharides

TOS transgalacto-oligosaccharides

XOS xylo-oligosaccharides

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