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PS Panesar and V Bali, Sant Longowal Institute of Engineering and Technology, Longowal, India
ã 2016 Elsevier Ltd. All rights reserved.
CH2OH
O OH
α O
HO
O OH OH
CH2OH O
O
O
HO
β
OH
n
OH O OH
CH2
n
O O
HO
CH2OH O HO HO
O O
OH OH
β
OH
CH2OH
Fructo-oligosaccharide Galacto-oligosaccharide
CH2OH
O
CH2OH
O CH2OH
HOH2C
CH2OH O
O OHHO O O O O
Gal1 O HO HO O
β 1
O HO HO O
4Gal1
β 4Gac1 1 OH OH HO CH2OH
OH OH
H.OH
OH
Xylo-oligosaccharide Soya-oligosaccharide
X OH OH H
OH N
O O O
H O OH
OH OH OH
OH OH OH
OH O O O
m n
Malto-oligosaccharide
OH OH
OH
CH2 OH OH
O
O
HO H OH
OH O OH
OH H OH O
H OH
O O O O
H OH
H OH
OH OH OH
OH
H OH HO
Lactulose Lactosucrose
extraction
Beet Raffinose
extraction
transglycosylation
Sucrose Fructo-oligosaccharides
transglycosylation
Cow’s milk Lactosucrose
extraction
isomerization
Lactose Lactulose
Starch transglycosylation
Galacto-oligosaccharides
hydrolysis
hydrolysis transglycosylation
Malto-oligosaccharides Glycosylsucrose
Soluble starch
hydrolysis
Isomalto-oligosaccharides
transglycosylation
extraction
Soybean extraction
Soybean whey Soybean oligosaccharides
hydrolysis
Xylo-oligosaccharides Xylan
hydrolysis
Chitin oligosaccharides Chitin
Figure 2 Production process of nondigestible oligosaccharides. Reproduced from Sako, T., Matsumoto, K. and Tanaka, R. (1999). Recent progress on
research and applications of non-digestible galacto-oligosaccharides. International Dairy Journal 9, 69–80, with permission.
that form 15–20% of starch possessing a-D-[1 ! 4] linkages) known as synbiotics, have associated health benefits. Prebi-
and amylo-pectin (highly branched and a major component otics have been studied for their role in reducing the risks of
possessing both a-D-[1 ! 4] and a-D-[1 ! 6) linkages]. Amylo- cancer, acute gastroenteritis, osteoporosis, cholesterol, and
pectin is a highly branched structure that provides more surface hyperlipidaemia; as well as for their bifidus-stimulating ability
area available for digestion in the small intestine. Resistant and immuno-modulatory effects.
starch possesses a higher amount of amylose, which limits
the exposed surface area. Therefore, starch with a high amylose
content is digested slowly, and some parts of it escapes diges-
tion in the small intestine and reaches colon. On the basis of its Production of Short Chain Fatty Acids
structure and complexity, resistant starch is further classified
The fermentative end products (short chain fatty acids) of pre-
into four classes. Type I includes physically inaccessible starch
biotic oligosaccharides produced by probiotic microorganisms
granules. Type II includes resistant starch granules. Type III
are in the order of acetate > propionate> butyrate in the ratio of
includes retrograded starch. Type IV includes chemically mod-
3:1:1 (Figure 3). Probiotic microorganisms use fermentation as
ified starches. Resistant starch naturally occurs in under-ripe
their energy source. Production of short chain fatty acids results
bananas, oats, lentils, white beans, etc. Commercially available
in a decreased pH, which leads to the reduced levels of patho-
resistant starch products include Hi-maize, CrystaLean, Nove-
genic bacteria, decreases bile acid solubility, and reduces ammo-
lose, Amylomaize VII, etc. These are reported to contribute in
nia absorption. Short chain fatty acids are water soluble and are
promoting apoptosis and reducing the risk of colon cancer.
readily absorbed into the blood stream by diffusion or through
anion exchange. They also influence intestinal motility. Only
5–10% of short chain fatty acids are eliminated in the feces.
The Prebiotic Effect Short chain fatty acids contribute to the growth and physiology
of intestinal tissues and systemic metabolism. Table 1 represents
Movement of food is rapid through the stomach and small
the function of various short chain fatty acids.
intestine (4–6 h) compared to colonic transit time, which is
around 48–70 h. This results in the formation of a stable
microbial community in the large intestine. Prebiotics (along
with other dietary fibers and undigested proteins) escape the Acute Gastroenteritis
upper part of the gastro-intestinal tract undigested, thus serving
as a potential substrate for probiotic endogenous bacteria Acute gastroenteritis is a disease state that occurs when food or
(bifidobacteria and lactobacilli) in the colon. Most of the water that is contaminated with pathogenic microorganisms
microflora in the colon are strict anaerobes and derive their (such as Clostridium perfringens, Vibrio cholera, E. coli,) or their
energy through fermentation. Prebiotics increase the popula- toxins is consumed. Some of its symptoms are nausea, vomit-
tion growth rate of bifidobacteria and lactobacilli from tenfold ing, diarrhea, and abdominal pain. Prebiotics help in overcom-
to a hundredfold. The presence of ‘osidic’ bonds makes pre- ing this problem by stimulating the growth of bifidobacteria
biotics nondigestible by human salivary and intestinal and lactobacilli, which act as a barrier against the invasion of
enzymes; therefore, they reach the colon in the undigested food pathogenic microorganisms. The probiotic bacteria com-
form. The hydrolysis and fermentation of these prebiotic oli- pete for nutrition and colonization sites, produce short chain
gosaccharides in the intestine by colonic microflora results in fatty acids, lower the pH, and secrete antimicrobial agents
their breakdown – mainly into short chain fatty acids. Pre- against the pathogenic bacteria, thereby reducing their inva-
biotics provide energy, metabolic substrate, and essential sion in the gut.
micronutrient selectively to the endogenous probiotic bacteria.
These probiotic bacteria have health related effects on the host
by reducing the risk of intestinal infections. The end product of
Cancer Risk
these prebiotics include short chain fatty acids (such as acetate,
butyrate, and propionate), lactate, pyruvate, ethanol, succi- Colorectal cancer has been reported mostly in people having a
nate, polyamines, bacitracin, and gasses such as H2, CO2, family history of colon cancer, high smoking rate, high body
CH4, H2S. Short chain fatty acids are mainly responsible for weight, greater consumption of alcohol and meat, lower phys-
causing the prebiotic effect. Short chain fatty acids lower lumi- ical workload, and lower intake of fruits and vegetables. Pre-
nal pH, which restricts the growth of disease-causing and biotics have been reported to enhance detoxification, reduce
putrefactive microorganisms like Escherichia coli, Salmonella, toxic metabolite production, and provide protection against
Listeria, and Shigella. Short chain fatty acids have also been tumor development in the colon. Prebiotics, along with pro-
reported to contribute in the regulation of absorption of biotics (bifidobacteria), have been studied to reduce a signifi-
sodium and water, maintenance of cholesterol, and increased cant number of colonic aberrant crypt foci and inhibit the
absorption of calcium and other minerals by the host. propagation phase of the carcinogen. The activity of colon
cancer causing bacterial enzymes such as b-glucuronidase,
b-glucosidase, and nitroreductase (which convert pre-
Health Benefits for Human carcinogens to proximal carcinogens) was observed to be
reduced by prebiotics by way of reducing the pH of the
Changes in dietary habits influence the balance of colonic colon. Prebiotics also reduce indole and isovaleric acids levels,
bacteria in the human gut. Probiotics and prebiotics, together which are markers of putrefaction.
Prebiotics 469
Fructans Xylans
Starch Pectins
Cellulose
Galactomannans
Arabinogalactan
Pentose
Glycolytic phosphate
pathway pathway
PEP-phosphoenolpyruvate
NADH NADPH
NAD+ NADP+
Formate
CO2 H2
CO2
Figure 3 The basic pathway for production of short chain fatty acid metabolites by bacterial fermentation in the colonic lumen. Reproduced
from Macfarlane, G. T. and Gibson, G. R. (1997). Carbohydrate fermentation, energy transduction and gas metabolism in the human large intestine.
In: Mackie, R. I. and White, B. A. (eds.) Ecology and physiology of gastrointestinal microbes: gastrointestinal fermentations and ecosystems,
pp. 269–318. New York: Chapman and Hall, with permission.
Table 1 Functions of short chain fatty acids minerals by modifying the electrolyte exchange, thus lowering
the risk of osteoporosis. Their effect on bones is further depen-
Short chain dent on the time, dose administered, and age of the host.
fatty acid Function
hypotriacylglycerolaemic effects. The increased production of microflora in the gut of animals and humans. Because of
short chain fatty acids through fermentation of prebiotics their properties, prebiotics (along with probiotics) can be
decreases, as well as inhibits, liver lipogenesis in hepatocytes. helpful in the treatment of various diseases such as acute
This may be due to modified gene regulation, which further gastroenteritis and osteoporosis. They can also reduce the risk
leads to down regulation of all hepatic lipogenic enzymes. of cancer, and help to lower cholesterol and hyperlipidaemia.
Prebiotics, like GOS, xylo-oligosaccharide, and isomalto- Prebiotics also possess the characteristics of having bifidus-
oligosaccharide, have been reported to have a significant role stimulating ability, immuno-modulatory effects, and low car-
in reducing serum cholesterol and triglycerides. iogenicity. These properties of prebiotics, along with their
safety aspects, have created countless new opportunities in
both the food industry, and in academia. With their low cost
Immune Modulation of production, the future is open for designing new prebiotic,
functional food products, as well as studying their structure,
Fermentative end products of prebiotics (i.e., short chain fatty
function, relationship, and their mode of action. Also, the
acids) have been reported to contribute to boosting the
increasing awareness of consumers is making prebiotics a
immune response by selective stimulation of specific mem-
promising market in the coming times.
branes of probiotic microflora. The G protein coupled receptors
of immune cells contain receptors that interact with short chain
fatty acids. These short chain fatty acids also play a significant
role in cytokine production, up regulation of gut-associated See also: Bifidobacteria in Foods: Health Effects; Colon: Structure and
lymphoid tissue expression of secretory IgA, anti-inflammatory Function; Dahi; Dairy Products: Dietary and Medical Importance; Fatty
profile, and increased chemotactic response of neutrophiles, Acids: Metabolism; Functional Foods; Lactic Acid Bacteria; Lactose;
thereby modulating the immune system of the host. Probiotics; Soy Beans: Dietary Importance.
Sajilata MG, Singhal RS, and Kulkarni PR (2006) Resistant starch – a review. Critical Swennen K, Courtin CM, and Delcour JA (2006) Non digestible oligosaccharides
Reviews in Food Science and Food Safety 5: 1–17. with prebiotic properties. Critical Reviews in Food Science and Nutrition
Sako T, Matsumoto K, and Tanaka R (1999) Recent progress on research and 46: 459–471.
applications of non-digestible galacto-oligosaccharides. International Dairy Journal Van Loo J, Coussement P, De Leenheer L, Hoebregs H, and Smits G (1995) Inulin and
9: 69–80. oligofructose in the western diets. Critical Reviews in Food Science and Nutrition
Stowell J (2007) Caloric control and weight management. In: Mitchell H (ed.) 35: 525–552.
Sweeteners and sugar alternatives in food technology. Oxford: Blackwell
Publishing Ltd.