168 CARBOHYDRATE METABOLISM
CARBOHYDRATE METABOLISM
R N Chibbar, S Ganeshan, M Båga, and
R L Khandelwal, University of Saskatchewan,
Saskatoon, SK, Canada
Published by Elsevier Ltd.
Introduction
Carbohydrates constitute two-thirds to three-quarters
of plant dry-matter, and are the major components of
plant storage organs. Cellulose, the most-abundant
plant carbohydrate provides structural integrity to
the plants and is used for fiber and fuel by humans.
In seeds, carbohydrates provide energy and substrates
for germination and initial growth of a new plant. The
major reserve carbohydrate present in grains is starch,
which constitutes a major source of calories in the
human diet and animal feed. Carbohydrates form
the basis of several important industries in the food
and feed sectors, and provide renewable and environ-
mentally friendly raw materials for industrial
applications such as biodegradable plastics, adhe-
sives, and ethanol-based fuels. With the development
of genetic modification technologies, the biosynthesis
of carbohydrates can be altered in planta to generate
novel products for various food, feed, and nonfood
applications. This article will describe the structure,
occurrence, and distribution of some of the common
storage and structural carbohydrates present in the
grains of major crops. The biochemical reactions oc-
curring during the biosynthesis of carbohydrates and
their utilization during seed germination will be
presented.
Carbohydrates
Definition and Classification
The term carbohydrate is derived from its basic ele-
mental formula [Cx(H2O)y], in which carbon is joined
to hydrogen and oxygen present in the same ratio as in
water. Chemically, carbohydrates are polyhydroxy
aldehydes or ketones, their simple derivatives or
their polymers. Carbohydrates in grains are classified
based on their chemical structures or their digestibility
when consumed by humans as food or by livestock as
feed. Based on the polymeric nature of carbohydrates,
they are classified as monosaccharides, disaccharides,
oligosaccharides, and polysaccharides (Figure 1).
Monosaccharides are the simplest of all sugars,
contain at least three carbon atoms, and are the
building blocks of all carbohydrates. The most
common monosaccharides in grains are the hexoses
(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,
the glycosidic bond is split to yield the component
monosaccharides. Sucrose and maltose are the most
commonly occurring disaccharides in most grains.
Oligosaccharides include the sugars that contain
3
20 sugar molecules joined by glycosidic bonds.
The most common oligosaccharides are soluble
a-galactosides, characterized by galactose moieties
joined by a(1,6) linkages. In legumes, raffinose, sta-
chyose, verbascose, and a-galactosides are the most
common oligosaccharides that constitute 6
18% of
the legume seeds dry weight. Polysaccharides are
polymers with more than 20 monosaccharides joined
together by glycosidic linkages. These are complex
molecules because of the diversity in monosaccharide
units and type of linkages present in the polysaccha-
rides. Some of the most common polysaccharides pre-
sent in grains are starch, cellulose, and xylans, of
which xylans include pentosans, b-glucans, and ara-
binoxylans. In relation to human and animal nutri-
tion, carbohydrates are also classified as available and
unavailable. The available carbohydrates represented
by sugars and starches are broken down and absorbed
by the human and animal digestive tract to give energy
to the body. Structural cell-wall carbohydrates such as
cellulose, hemicellulose, and nonstarch complex car-
bohydrates, e.g., xylans are categorized as unavailable
carbohydrates because they are not assimilated by the
digestive tract. The unavailable carbohydrates are
also often referred to as dietary fibers.
Distribution in Important Grains
All the above-described carbohydrates are present in
most grains, where they form the major constituent of
the dry weight (Table 1). However, in oil seed grains,
such as Brassica spp., lipids and proteins are the major
storage grain components and carbohydrates make up
only one-quarter of the grain dry-weight. In cereal
grains, the total concentration of mono-, di-, and oli-
gosaccharides varies from less than 1
3% (Table 2).
Corn kernels contain the highest percentage of glucose
and fructose, whereas in barley, sucrose is the major
free sugar. Wheat grains have the highest concentra-
tion of raffinose among the cereal grains (Table 2).