4.

ESSENTIAL NUTRIENTS - CARBOHYDRATES

4.1 Carbohydrates
After the proteins and lipids, the carbohydrates represent the third most abundant group of
organic compounds in the animal body. By contrast, carbohydrates constitute the major class of
organic nutrients within plant tissues. The carbohydrate group includes such important
compounds as glucose, fructose, sucrose, lactose, starch, glycogen, chitin, and cellulose.
Carbohydrates are usually defined as substances containing carbon, hydrogen and oxygen,
with the last two elements being present in the same ratio as in water (ie. Cx(H2O)y ). Although
this definition is satisfactory for the majority of compounds present within this group, a few
carbohydrates contain a lower proportion of oxygen than that in water or exist as carbohydrate
derivatives which may contain nitrogen and sulphur.
4.1.1 Classification
The carbohydrates can be divided into two major groups according to their chemical structure;
the sugars and non-sugars (Table 8). The simplest sugars are called monosaccharides, and
these inturn can be divided into five sub-groups depending on the number of carbon atoms
present in the molecule: Trioses (C3H6O3), Tetroses (C4H8O4), Pentoses (C5H10O5), and Hexoses
(C6H12O6) These monosaccharides may also inturn be linked together (with the elimination of
water) to form di, tri or polysaccharides containing two, three or more monosaccharide units or
residues respectively. Here the term ‘sugar’ is restricted to those carbohydrates containing less
than 10 monosaccharide units. Non-sugars are therefore carbohydrates which contain more
than 10 monosaccharide units and which do not possess a sweet taste. The non-sugars can be
divided into two sub-groups, homopolysaccharides and heteropolysaccharides; the former
consisting of identical monosaccharide units and the latter of mixtures of different
monosaccharide units.
4.1.2 Monosaccharides
In general all monosaccharides are soluble in water, sparingly soluble in ethanol and insoluble
in ether, are optically active, possess reducing properties (ie. reduce Fehling's solution), can be
represented by the general formula CxH2xOx, and generally possess a sweet taste.
The chain formula of some of the more common monosaccharides can be represented thus;
Triose (C3H6O3) Pentose (C5H10O5) Hexose (C6H12O6) Hexose (C6H12O6)

D(+)-glyceraldehyde D(+)-ribose D(+)-glucose D(+)-galactose

Table 8. Carbohydrate classification
Hexose (C6H12O6) Hexose (C6H12O6)
D(+)-mannose D(-)-fructose
Within the structural formulae presented ‘D’ represents the configuration or direction of the
hydroxyl group (OH) on the carbon atom next to the last carbon from the functional or aldehyde
group. For example, in the case of D(+)-glyceraldehyde and D(+)-glucose the hydroxyl group on
the penultimate carbon atom (ie. C2 and C5 respectively) is on the right when the aldehyde
group (CHO) is at the top of the formula. Similarly, the symbol (+) or (-) denotes the direction of
optical rotation observed when a solution of that sugar is placed in a polarimeter; dextro-rotatory
(clockwise direction, +) or laevorotatory (anti-clockwise direction, -). Virtually all naturally
occuring monosaccharides are members of the D series of sugars, the configuration around the
penultimate carbon atom being the same as that in D-glyceraldehyde. In addition, all naturally
occuring monosaccharides are dextro-rotatory, with the exception of fructose and erythrose.
There is considerable evidence to suggest that monosaccharides may also exist in a ring or
cyclic molecular form. For example, two cyclic forms of D-glucose are known to occur in nature;
α-D-glucose and β-D-glucose. As with the chain formulae, the difference between these two
cyclic forms depends upon the configuration or direction of the hydroxyl group on carbon atom
1.

α-D-glucose β-D-glucose
The biological importance of the structural difference between α and β -D-glucose must be
stressed here; the structural configuration determining the physical and subsequent biological
properties of polysaccharides composed of individual monosaccharide units. For example, the
polysaccharide cellulose is composed of insoluble zig-zag chains of β-glucose units, whereas
the polysaccharides starch and glycogen are composed of more biological reactive helical or
branched chains of α-glucose units.
It should be mentioned at this stage that monosaccharides are seldom directly involved in
biochemical reactions within the cell, but are first transformed into appropriate monosaccharide
derivatives. Important monosaccharide derivatives include the sugar phosphate esters (D-
glucose-6-phosphate, D-glucose-1-phosphate, D-fructose-6-phosphate, and phosphate
diesters), amino sugars (D-glucosamine), sugar acids (gluconic acid, glucuronic acid) and sugar
alcohols (sorbitol).

D-glucose-6-phosphate β-glucuronic acid α-D-glucosamine

Pentoses
Important pentose monosaccharides include L-arabinose, D-xylose and D-ribose. From a
nutritional viewpoint the most important pentose is D-ribose and its derivatives D-deoxyribose
and ribitol. For example, D-ribose and D-deoxyribose are essential components of ribonucleic
acid (RNA) and deoxyribose nucleic acid (DNA) respectively, and ribitol an essential component
of the vitamin riboflavin.

α-D-ribose α-D-deoxyribose
Hexoses
Glucose: occurs in its free form in plant tissues, fruits, honey and blood. In most natural
foodstuffs glucose is present in a combined form, either as the sole monosaccharide component
of disaccharides (ie. maltose) and polysaccharides (ie. starch, glycogen, cellulose), or combined
with other monosaccharides in the form of lactose (milk sugar), sucrose and
heteropolysaccharides. Glucose is fermented by yeast during the manufacture of beers and
wine to yield alcohol and carbon dioxide. Similarly, as fruit ripen their free sugar or glucose
content normally increases and starch content decreases accordingly.
Fructose: like glucose, fructose is found in its free state in plant juices, fruit and honey. It is a
componet of the disaccharide sucrose, and is the sweetest sugar known in nature (ie. it is
responsible for the exceptional sweet taste of honey).
Galactose: although galactose does not occur in its free state in nature, it exists as a component
of the disaccharide lactose and many polysaccharides, including galactolipids, gums, and
mucilages.
4.1.3 Disaccharides
Disaccharides are composed of two hexose sugars joined together with the elimination of water,
thus;
C6H12O6 + C6H12O6 = C12H22O12 + H2O
The most important naturally occuring disaccharides are maltose, sucrose and lactose.
Maltose: is composed of two molecules of glucose joined together through a α-1,4-glycoside
linkage. Maltose is a reducing sugar and is soluble in water.

Maltose is not found in nature to any extent but is a product obtained during the enzymatic
degradation of starch. For example, maltose is produced from starch during the germination of
barley by the action of the enzyme amylase; after germination and drying the barley (or ‘malt’ as
it is now called) is used in the manufacture of beer and malt whisky.
Sucrose: is composed of one molecule of glucose and fructose joined together through aα-1, β-
2-glycoside linkage. Since both functional reducing groups are involved in the glycoside linkage,
sucrose does not possess reducing properties.

Sucrose is widely distributed in nature and occurs in most plants; rich sources of sucrose
include sugar cane (20% sucrose), sugar beet (15–20%), mangolds and carrots. Sucrose is the
sugar of familiar use in the domestic household. When sucrose is heated to a temperature of
160°C it forms barley sugar, and at 200°C forms caramel. Beet or cane molasses are
agricultural by-products obtained from the manufacture of sucrose from sugar beets or sugar
cane respectively. Molasses are dark coloured, viscous liquids (20–30% moisture), from which
no further sucrose can be separated by usual crystallization procedures because of the
presence of appreciable quantities of reducing sugars (ie. glucose) and non-sugar impurities.
Lactose: is composed of one molecule of glucose and galactose joined together by an β-1,4-
glycoside linkage. Like maltose it has reducing properties. Lactose, or milk sugar, is the
principal sugar found in milk and is unique to mammals. It forms about 40% of the total milk
solids; the total lactose content of cows milk and human milk being 4.6–4.8% and 7%
respectively. Lactose readily undergoes bacterial fermentation; for example the souring of milk
by Streptococcus lactis by the fermentation of lactose to lactic acid. Like sucrose, if lactose is
heated to a temperature of 175°C it forms lactocaramel.

4.1.4 Homopolysaccharides
These carbohydrates are very different from the sugars. They have a high molecular weight and
are composed of large numbers of hexose or to a lesser extent pentose residues. Many of them
occur in plants or animals as a reserve food material (ie. starch or glycogen) or as structural
materials (ie. cellulose or chitin).
Starch: is composed of two structural components, amylose and amylopectin. Although the
relative proportions of amylose and amylopectin within plant starches varies depending on the
species (20–30% amylose and 70–80% amylopectin), the fundamental unit of these two
structural components is α-D-glucose. For example, amylose consists of long unbranched
chains of 100 or more D-glucose units joined together by α-1,4 linkages. On the otherhand,
amylopectin is composed of highly branched chains of D-glucose units (20–30 units per
branch); the units being joined together by α-1,4 linkages and also α-1,6 linkages (the α-1,6
glycoside linkages used only at the beginning of a side chain).
Starch is the depot form of sugar or glucose in plants; it occurs in stems, fruits, seeds, roots or
leaves, forming the greatest carbohydrate food reserve of plants and consequently constitutes
the largest carbohydrate component of animal feeds. For example, starch may account for up to
70% by weight of seeds and up to 30% of fruit, tubers, or roots. Starch is stored within plants in
the form of granules, the size and shape of which varies from species to species. Each granule
is surrounded by a thin layer of cellulose which render them insoluble in water and undigestible
in their raw or uncooked form by non-ruminant animals including fish and shrimp. Cooking, by
heating in the presence of moisture will however facilitate the rupture of the cellulose
membrane, resulting in the absorption of water by the starch within, which in the presence of
heat will result in the gelatinization of the starch forming a gelatinous solution or paste. When
starch is subjected to dry heat, dextrin is formed; dextrin being an intermediate degradation
product of starch in the sequence starch → dextrin → maltose → glucose. For example, starch
in bread is converted to dextrin when toasted, the dextrin giving toast its characteristic taste.
Glycogen: is composed of branched chains of α-D-glucose units joined together by α -1,4
linkages and α -1,6 linkages; the latter being more numerous in glycogen (as compared with
amylopectin) due to the presence of more and shorter branches of 10–20 glucose units.
Glycogen is the form in which carbohydrate is stored within the animal body; being particularly
concentrated in the liver and muscle.
Cellulose: is composed of very long chains of D-glucose units joined together by β-1,4 linkages.
It is a very stable polysaccharide and is the most abundant carbohydrate in nature; forming the
fundamental structure of the plant cell wall. Cellulose has great tensile strength and is resistant
to chemical attack. Although cellulose can be hydrolysed by strong acid treatment, with the
exception of micro-organisms, few non-ruminant animals have the necessary endogenous
enzymes (ie. cellulases) capable of hydrolysing and digesting cellulose. Forexample, the
cellulase enzymes which are capable of attacking cellulose are only found in germinating seeds,
fungi and bacteria (ie. such as those present in the digestive tract of ruminants). An example of
a nearly pure form of cellulose is cotton.
Chitin: is composed of repeating units of N-acetyl-D-glucosamine joined together by β -1,4
linkages and is therefore similar in structure to cellulose.

Chitin is the major structural component of the cuticle of insects and the exoskeleton of
crustaceans.
4.1.5 Heteropolysaccharides
In contrast to the homopolysaccharides, the heteropolysaccharides consist of high molecular
weight mixtures of different monosaccharide units.
Hemicellulose: is composed principally of xylose units linked together by β -1,4 linkages, but
may also contain hexoses and sugar acids (ie. uronic acid). These polysaccharides normally
accompany cellulose in the leafy and woody structure of higher plants and seeds. They are
insoluble in water and like cellulose are not easily digested by non-ruminant animals.
Gums: are constituents of plant wounds and are very complex compounds; on hydrolysis
yielding a variety of monosaccharide and sugar acids. An example of a gum is gum arabic
(acacia gum).
Mucilages: complex carbohydrates present in certain plants and seeds. Many algae, and
especially seaweeds, yield mucilages which are soluble in hot water and which form a gel on
cooling. Agar, a sulphuric acid polymer of galactose, is a widely used mucilage or gel obtained
from red seaweeds (Gelidium family). Other examples include alginic acid derived from brown
seaweeds (Laminaria family).
Pectic substances: complex carbohydrates which contain D-galacto-uronic acid as the main
constituent. They occur mainly in the primary cell wall and intercellular layers of land plants, and
are particularly abundant in citrus fruit, sugar beet, apples, and some root vegetables (ie.
turnips). As with mucilages, pectic acids have strong gelling properties and are often used in the
preparation of jams.
Mucopolysaccharides: complex carbohydrates which contain amino sugars and uronic acids,
and constitute the mucous secretions of animals. They are acidic in nature and may be rich in
sulphate ester groups; important mucopolysaccharides include chondroitin sulphate (present in
cartilage, bone, heart valves, tendons and the cornea of the eye), heparin (anti-coagulant
present in blood vessels, liver, lung, and spleen), and hyaluronic acid (viscous lubricating fluid
present in the skin, vitreous humour of the eye, the synovial fluid of joints, and the umbilical cord
of mammals). Finally, mucopolysaccharides constitute the principal component of the cell wall of
many bacteria; the mucopolysaccharide cell wall consisting of alternating units of N-acetyl-
muramic acid and N-acetyl-glucosamine linked to short peptide chains.
4.2 Carbohydrate function
Carbohydrates are synthesized in all green plants by a process called photosynthesis, which
can be represented thus;
6CO2 + 6H2O + Light  → C6H12O6 + 6O2
  (673 kcal)
Within man and terrestrial farm animals dietary carbohydrates serve as the principal source of
metabolic energy (ATP). This reaction can be represented as;
C6H12O6 + 6O2   →   6CO2 + 6H2O + 38 ATP1
In fish and shrimp no absolute dietary requirement for carbohydrate has been established to
date. This contrasts markedly with that of dietary protein and lipid, where specific dietary
requirements have been established for certain essential amino acids and fatty acids. To a large
extent this has been due:
 The carnivorous/omnivorous feeding habit of the majority of farmed fish and and shrimp
species.
 The ability of fish and shrimp to synthesize carbohydrates (ie. glucose) from non-
carbohydrate substrates such as protein and lipid (a process called gluconeogenesis).
 The ability of fish and shrimp to satisfy their dietary energy requirements through protein
and lipid catabolism alone if so required.
However, despite the apparent absence of a dietary requirement for carbohydrate in fish or
shrimp, there is no doubt that carbohydrates perform many important biological functions within
the animal body. For example; glucose, the end product of carbohydrate digestion in animals,
serves as the major energy source of brain and nervous tissue, and as a metabolic intermediate
for the synthesis of many biologically important compounds, including the chitin exoskeleton of
crustacea, the nucleic acids RNA and DNA, and the mucopolysaccharide mucous secretions.
Although carbohydrates may be regarded as non-essential dietary nutrients for fish and shrimp,
their inclusion in practical diets is warranted because:
 They represent an inexpensive source of valuable dietary energy for noncarnivorous fish
and shrimp species.
 Their careful use in practical diets can spare the more valuable protein for growth
instead of energy provision (a procedure called ‘protein sparing’.
 They serve as essential dietary constituents for the manufacture of water stable diets
when used as binders (ie. gelatinized starch, alginates, gums).
 Certain carbohydrate sources serve as dietary components which can increase feed
palatability and reduce the dust content of finished feeds (ie. cane or beet molasses).
1
ATP or Adenosine triphosphate contains ‘high energy’ bonds, and is the the form in which a large part of the liberated energy is
stored within the body.

4.3 Carbohydrate metabolism
Although glycogen constitutes the major fuel source during anaerobic metabolism (glycolysis)
within the white muscle of fish during ‘burst’ swimming activities, the ability of the liver and
tissues to store glycogen is limited; total carbohydrates as glycogen constituting less than one
percent of the wet body tissue (Cowey and Sargent, 1979). By contrast, juvenile shrimp P.
japonicus have been found to contain glucose, acetyl glucosamine and trehalose as the major
carbohydrates of body tissue (Kanazawa, 1983).
In contrast to omnivorous mammals, fish do not mobilize liver glycogen rapidly when they are
starved. In fact it has been shown within starving fish that the oxidation of non-carbohydrate
substrates takes precedence over the mobilization and hydrolysis of glycogen. This suggests
that the capacity of fish to oxidize glucose aerobically is somewhat limited. It follows therefore
that gluconeogenesis may play a major role in maintaining blood sugar levels in starving or
fasting fish. For example, for cultured fish fed a high protein diet it is probable that the energy
demand of tissues (ie. brain and nervous tissue) which catabolise glucose are met by
gluconeogenesis (from amino acids and triacylglycerol) rather than by glycogenolysis (for review
see Walton and Cowey, 1982). However, studies with the eel (A. japonica) would appear to
contradict the above general hypothesis (Degani, Viola and Levanon, 1986).
4.4 Dietary carbohydrate utilization
The ability of carnivorous fish species to hydrolyze or digest complex carbohydrates is limited
due to the weak amylotic activity in their digestive tract (Spannhof and Plantikow, 1983). Thus,
for fish species such as trout, as the proportion of dietary starch is increased, starch digestibility
decreases accordingly (Singh and Nose, 1967; Bergot and Breque, 1983). Furthermore, in long
term feeding trials with carnivorous fish species (ie. salmonids) it has been shown that high
dietary carbohydrate levels depress growth, elevate liver glycogen levels, and cause eventual
mortality (Phillips et. al., 1948; Austreng et al., 1977). By contrast, warmwater omnivorous or
herbivorous fish speceis such as carp (C. carpio), channel catfish (I. punctatus), tilapia (O.
niloticus), and eel (A. japonica) have been found to be more tolerant of high dietary
carbohydrate levels; the dietary carbohydrate being effectively utilized as a dietary energy
source or excess stored in the form of body lipid (Chiou and Ogino, 1975; Robinson and Wilson,
1985; Anderson et al., 1984; Degani, Viola and Levanon, 1986).
The utilization of dietary carbohydrate has also been found to vary with the complexity or
chemical structure of the carbohydrate source used (digestible polysaccharides and
disaccharides having a more beneficial effect on growth than monosaccharides: fish - Pieper
and Pfeffer, 1980; Robinson and Wilson, 1985; Anderson et al., 1984; shrimp - Alava and
Pascual, 1984; Deshimaru, 1981; 1 Kanazawa, 1983), the physical state of the carbohydrate
source used (cooked or gelatinized starches having a higher digestibility and beneficial effect on
growth than native or raw starches: fish - Spannhof and Plantikow, 1983; Bergot and Breque,
1983; Robinson and Lovell, 1984), and daily feed intake (a restricted feeding regime having a
beneficial effect on starch digestibility: fish - Bergot and Breque, 1983). From the above
discussion it appears that the ability of fish or shrimp to adapt to high carbohydrate diets
depends on their ability to convert excess energy (ie. glucose) into lipids or non-essential amino
acids.
1
Furuichi, Taira and Yone (1986): availability of glucose in yellowtail (S. quinqueradiata) is lower than that of alpha starch.
Since most farmed fish species have a relatively short gastro-intestinal tract that does not lend
its self to the development of an extensive bacterial flora (as in a ruminant animal), the intestinal
cellulase activity of fish from resident bacteria is weak or absent (Stickney and Shumway,
1974). It follows therefore that dietary cellulose or ‘crude fibre’ (ie. dietary carbohydrates which
are resistant to chemical treatment with dilute acid or alkali, including cellulose and
hemicellulose) has no utilizable energy value to fish, and in dietary excess has a deleterious
effect on growth and feed efficiency (Anderson et al., 1984; Poston, 1986; Hilton, Atkinson and
Slinger, 1983; Bromley and Adkins, 1984).