Carbohydrates represent a broad group of substances which include the sugars, starches, gums and celluloses.

The common attributes of carbohydrates are that they contain only the elements carbon, hydrogen and oxygen,
and that their combustion will yield carbon dioxide plus one or more molecules of Water.

The simplest carbohydrates are the three-carbon sugars which figure importantly in intermediary metabolism
and the most complex are the naturally occurring polysaccharides, primarily of plant, origin. In the diet of
animals and fish, two classes of polysaccharides are significant:
(a) structural polysaccharides which are digestible by herbivorous species -cellulose, lignin, dextrans, mannans,
inulin, pentosans, pectic acids, algic acids, agar and chitin; and
(b) universally digestible polysaccharides - principally starch.
Carbohydrates make up three-fourths of the biomass of plants but are present only in small quantities in the
animal body as glycogen, sugars and their derivatives. Glycogen is often referred to as animal starch because it
is not present in plants. Derived mono-saccharides such as the sugar acids, amino sugars and the deoxysugars
are constituents of all living organisms.
Carbohydrates, one of the four major macronutrients, provide a significant amount of fuel to the human body.
However, if carbohydrates are not properly digested and absorbed, they cannot perform their essential functions.
Digestion and absorption occurs along the gastrointestinal tract, and remaining, undigested carbohydrates are
then eliminated from the colon.
Types of Carbohydrates
There are three main types of carbohydrates: starches, sugars and dietary fiber.
Starches and sugars are considered the energy-yielding carbohydrates because they are fully digestible and, once
absorbed, they provide the body with 4 calories of energy per gram of carbohydrate.
Alternatively, fiber is a type of carbohydrate which is not fully digestible because humans lack the enzymes to
break down fibers. As such, fiber is the main carbohydrate which is eliminated through excretion.

CLASSIFICATION AND CHEMISTRY

Carbohydrates are classified generally according to their degree of complexity.
Hence, the free sugars such as glucose and fructose are termed monosaccharides;
sucrose and maltose, disaccharides;
and the starches and celluloses, polysaccharides.
Carbohydrates of short chain lengths such as raffinose, stachyose and verbascose, which are three, four and five
sugar polymers respectively, are classified as oligosaccharides.
2.1 Pentoses
Pentoses are five-carbon sugars seldom found in the free state in nature. In plants they occur in polymeric forms
and are collectively known as pentosans. Thus, xylose and arabinose are the constituents of pentosans present in
plant fibres and vegetable gums, respectively. As the sugar moieties in nucleic acids and riboflavin, ribose and
deoxyribose are indispensable constituents of the life process. D-ribose has the following chemical structure:
CHO
Ι
H -C-OH
Ι
H -C-OH
Ι
H -C-OH
Ι
CH2-OH

D-Ribose
2.2 Hexoses
The hexoses comprise a large group of sugars. Principal among these are: glucose, fructose, galactose and
mannose. While glucose and fructose are found free in nature, galactose and mannose occur only in combined
form. The hexoses are divided into aldoses and ketoses according to whether they possess aldehydic or ketonic
groups. Thus, glucose is an aldo sugar and fructose is a keto sugar. The presence of aymmetric centres in all
sugars with three or more carbon atoms gives rise to stereoisomers. Galactose and mannose are stereoisomers of
glucose which, theoretically, is only one of 16 stereoisomers. Because the ketohexoses have only three
asymmetric centres, fructose is one of eight stereoisomers. The chemical configurations of the four hexoses
mentioned are as follows:

D-Glucose D-Galactose D-Mannose D-Fructose

A general phenomenon, known as mutarotation, is observed in a variety of pentoses and hexoses as well as in
certain disaccharides. For example, it has been established that two isomers of D-glucose exist, hence requiring
an additional asymmetric centre in this sugar. It became apparent that D-glucose and most other sugars have
cyclic structures. The position of the hydroxyl group in relation to the ring oxygen characterizes this additional
configurations modification. By convention, the positioning of the hydroxyl group on carbon atom 1 on the
same side of the structure as the oxygen ring describes a -modification; and, the positioning of the same
hydroxyl group on the opposite side of the ring oxygen describes a b -modification.

a -D-Glucose b -D-Glucose
Carbohydrases, which catalyse the hydrolysis of glycosidic linkages of simple glycosides, oligosaccharides and
polysaccharides often exhibit specificity with regard to substrate configuration. As we shall see later, the
specificity for enzyme hydrolysis of certain oligosaccharides helps to explain the poor utilization of this class of
carbohydrates in fish nutrition.
Sugars containing the aldo or the keto group are capable of reducing copper in alkaline solutions (Fehling's
solution) to produce the brick-red colouration of cuprous ions. These sugars are called reducing sugars and the
reaction, although not specific for reducing sugars, has use for both qualitative and quantitative determinations.
Glucose is widely distributed in small amounts in fruits, plant juices and honey. It is commercially produced by
the acid or enzyme hydrolysis of grain and root starches. Glucose is of special interest in nutrition because it is
the end-product of carbohydrate digestion in all non-ruminant animals including fish.
Fructose is the only important ketohexose and is found in the free state alongside glucose in ripening fruits and
honey. Combined with glucose it forms sucrose. Fructose is somewhat sweeter than sucrose and is produced in
increasing quantities commercially as a sweetener.
Galactose occurs in milk in combination with glucose. It is also present in oligo-saccharides of plant origin, in
combination with both glucose and fructose.
Mannose is present in some plant polysaccharides collectively termed mannans.
2.3 Disaccharides
Disaccharides are condensation products of two molecules of monosaccharides. Sucrose is the predominant
disaccharide occurring in the free form and is the principal substance of sugar cane and sugar beet. It is also
formed during germination of legume seeds. Other common disaccharides are maltose and lactose. Maltose is a
dimer of glucose, and lactose is a copolymer of galactose and glucose. The two molecules of glucose in maltose
are held together in an a -1,4 glycosidic linkage whereas the two hexose entities of galactose are linked at the b -
1,4 position. Glucose and fructose are combined in an a -1,2 linkage in sucrose. The abbreviated name of
sucrose is D-Glu-(a, 1® 2)-D-Fru.

a -Maltose
b -Lactose

Sucrose

2.4 Oligosaccharides
The Oligosaccharides raffinose, stachyose and verbascose are present in significant quantities in legume seeds.
Raffinose, which is the most widespread among the three, consists of one molecule of glucose linked to a
molecule of sucrose at the a -1,6 position. Its abbreviated chemical name is a -D-Gal (1® 6) -a - D -Glu - (1® 2)
- b -D-Fru. Further chain elongation at the galactose end with another galactose molecule will yield stachyose.
These galactose-galactose linkages are all at the a-l,6 position and digestion of these Oligosaccharides by
animals requires a highly specific enzyme not elaborated by the animals themselves but by certain bacteria
present in the animals guts. The gradual disappearance of oligosaccharides from the cotelydons of legume seeds
during germination is part of an intricate process beginning with uptake of water by the seed. This uptake of
moisture releases gibberellic acid which in turn activates the DNA in the seed, thereby triggering the life cycle
of the plant. The DNA directs the production of a -galactosidase which is required for the hydrolysis of these
Oligosaccharides. Any interference of the DNA transcription process blocks enzyme production and will be
evidenced by continued senescence of the seed and persistence of oligosaccharides in the seed cotelydons.
2.5 Polysaccharides
The polysaccharides represent a large group of complex carbohydrates which are condensation products of
undetermined numbers of sugar molecules. The various subgroups are rather ill-defined and there is a lack of
agreement on their classification. Most polysaccharides are insoluble in water. Upon hydrolysis with acids or
enzymes they eventually yield their constituent monosaccharides.
Starch is a high molecular weight polymer of D-glucose and is the principal reserve carbohydrate in plants.
Most starches consist of a mixture of two types of polymers, namely; amylose and amylopectin. The proportion
of amylose and amylopectin is generally one part of amylose and three parts of amylopectin. Enzymes capable
of catalyzing the hydrolysis of starch are present in the digestive secretions of animals and fish within their
cells. The a-amylases which are found virtually in all living cells cleave the a -D-(1® 4) linkages at random and
bring about an eventual total conversion of the starch molecule into the reducing sugars. The principal  a -
amylases of animal origin are those produced in the salivary gland and the pancreas. Starch is insoluble in water
and is stained blue by iodine.
Glycogen is the only complex carbohydrate of animal origin. It exists in limited quantities in liver and muscle
tissues and acts as a readily available energy source.
Dextrins are intermediate compounds resulting from incomplete hydrolysis or digestion of starch. The presence
of a -D-(1® 6) linkages in amylopectin and the inability of a -amylase to cleave these bonds give rise to low
molecular weight carbohydrate segments called limit dextrins. These residues are acted upon primarily by
acidophilic bacteria in the digestive tract.
Cellulose is made up of long chains of glucose units held together by b -D-(1® 4) linkages. The enzymes which
cleave these linkages are not ordinarily present in the digestive secretions of animals and fish although some
species of shellfish are believed to elaborate cellulase, the enzyme which catalyzes the hydrolysis of cellulose.
Cellulase producing micro-organisms present in the gut of herbivorous animals and fish impart to their host
animals the ability to utilize as food the otherwise indigestible cellulose.
Other complex polysaccharides in common occurrence are the hemicelluloses and pentosans. Hemicellulose
represents a group of carbohydrates including araban, xylan, certain hexosans and polyuronides. These
substances are generally less resistant to chemical treatment and undergo some degree of enzymatic hydrolysis
during normal digestive processes. Pentosans are polymers of either xylose or arabinose as constituents of plant
structural material and vegetable gums, respectively.

Carbohdyrate Digestion
The two digestible carbohydrates are starches and sugars, and both of these carbohydrates are digested, or
broken down into their most elementary form, along the gastrointestinal tract. Amylase, an enzyme which
breaks apart starches, is found in the mouth and in the small intestine. Similarly, the three major enzymes which
break apart sugars -- sucrase, maltase and lactase -- are also found in the mouth and in the small intestine. Once
these digested starches and sugars begin to move through the small intestine, they are able to be absorbed.
Carbohydrate Absorption
Once carbohydrates are broken down into their simplest forms, they are quickly absorbed along the upper and
lower parts of the small intestine. Small, finger-like projections, called villi, absorb the carbohydrates, then they
are transferred to the blood stream and carried to muscles and the liver.
Carbohydrate Elimination
When carbohydrates are not fully digested or absorbed, they are eliminated from the body. Dietary fiber is one
of the carbohydrates which humans cannot digest, thus dietary fiber is the most commonly excreted type of
carbohydrate. In addition, lactose, a type of sugar, can also be excreted if an individual lacks the proper enzymes
to digest this carbohydrate. All undigested carbohydrates move from the small intestine, where absorption
would normally occur, to the large intestine and the colon, where elimination finally occurs.