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168 CARBOHYDRATE METABOLISM

CARBOHYDRATE METABOLISM

R N Chibbar, S Ganeshan, M Båga, and contain at least three carbon atoms, and are the
R L Khandelwal, University of Saskatchewan, building blocks of all carbohydrates. The most
Saskatoon, SK, Canada common monosaccharides in grains are the hexoses
Published by Elsevier Ltd. (fructose, glucose, and galactose) and the pentoses
(arabinose and xylose). Two sugar moieties joined
by a glycosidic bond (an oxygen bridge) form
a disaccharide. Upon hydrolysis of disaccharides,
Introduction the glycosidic bond is split to yield the component
monosaccharides. Sucrose and maltose are the most
Carbohydrates constitute two-thirds to three-quarters commonly occurring disaccharides in most grains.
of plant dry-matter, and are the major components of Oligosaccharides include the sugars that contain
plant storage organs. Cellulose, the most-abundant 320 sugar molecules joined by glycosidic bonds.
plant carbohydrate provides structural integrity to The most common oligosaccharides are soluble
the plants and is used for fiber and fuel by humans. a-galactosides, characterized by galactose moieties
In seeds, carbohydrates provide energy and substrates joined by a(1,6) linkages. In legumes, raffinose, sta-
for germination and initial growth of a new plant. The chyose, verbascose, and a-galactosides are the most
major reserve carbohydrate present in grains is starch, common oligosaccharides that constitute 618% of
which constitutes a major source of calories in the the legume seeds dry weight. Polysaccharides are
human diet and animal feed. Carbohydrates form polymers with more than 20 monosaccharides joined
the basis of several important industries in the food together by glycosidic linkages. These are complex
and feed sectors, and provide renewable and environ- molecules because of the diversity in monosaccharide
mentally friendly raw materials for industrial units and type of linkages present in the polysaccha-
applications such as biodegradable plastics, adhe- rides. Some of the most common polysaccharides pre-
sives, and ethanol-based fuels. With the development sent in grains are starch, cellulose, and xylans, of
of genetic modification technologies, the biosynthesis which xylans include pentosans, b-glucans, and ara-
of carbohydrates can be altered in planta to generate binoxylans. In relation to human and animal nutri-
novel products for various food, feed, and nonfood tion, carbohydrates are also classified as available and
applications. This article will describe the structure, unavailable. The available carbohydrates represented
occurrence, and distribution of some of the common by sugars and starches are broken down and absorbed
storage and structural carbohydrates present in the by the human and animal digestive tract to give energy
grains of major crops. The biochemical reactions oc- to the body. Structural cell-wall carbohydrates such as
curring during the biosynthesis of carbohydrates and cellulose, hemicellulose, and nonstarch complex car-
their utilization during seed germination will be bohydrates, e.g., xylans are categorized as unavailable
presented. carbohydrates because they are not assimilated by the
digestive tract. The unavailable carbohydrates are
Carbohydrates also often referred to as dietary fibers.

Definition and Classification Distribution in Important Grains


The term carbohydrate is derived from its basic ele- All the above-described carbohydrates are present in
mental formula [Cx(H2O)y], in which carbon is joined most grains, where they form the major constituent of
to hydrogen and oxygen present in the same ratio as in the dry weight (Table 1). However, in oil seed grains,
water. Chemically, carbohydrates are polyhydroxy such as Brassica spp., lipids and proteins are the major
aldehydes or ketones, their simple derivatives or storage grain components and carbohydrates make up
their polymers. Carbohydrates in grains are classified only one-quarter of the grain dry-weight. In cereal
based on their chemical structures or their digestibility grains, the total concentration of mono-, di-, and oli-
when consumed by humans as food or by livestock as gosaccharides varies from less than 13% (Table 2).
feed. Based on the polymeric nature of carbohydrates, Corn kernels contain the highest percentage of glucose
they are classified as monosaccharides, disaccharides, and fructose, whereas in barley, sucrose is the major
oligosaccharides, and polysaccharides (Figure 1). free sugar. Wheat grains have the highest concentra-
Monosaccharides are the simplest of all sugars, tion of raffinose among the cereal grains (Table 2).

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