Carbohydrates

All carbohydrates contain the elements carbon, hydrogen and oxygen. The ‘hydrate’ part of the
name comes from the fact that hydrogen and oxygen atoms are present in the ratio of 2 : 1, as they
are in water (‘hydrate’ refers to water).

The general formula for a carbohydrate can therefore be written as Cx(H2O)y. Carbohydrates are
divided into three main groups, namely monosaccharides, disaccharides and polysaccharides. The
word ‘saccharide’ refers to a sugar or sweet substance.

Monosaccharides
Monosaccharides are sweet-tasting, soluble substances that have the general formula (CH2O)n.
While “n” can be any number from 3 to 7. Monosaccharides consist of a single sugar molecule
(‘mono’ means one). The main types of monosaccharides, if they are classified according to the
number of carbon atoms in each molecule, are trioses (3C), pentoses (5C) and hexoses (6C). The
names of all sugars end with -ose. Common hexoses are glucose, fructose and galactose. Two
common pentoses are ribose and deoxyribose.

Glucose is the best-known monosaccharide. This molecule is a hexose and has a formula of
C6H12O6. However, the atoms of carbon, hydrogen and oxygen can be arranged in many different
ways. These atoms form a ring which can take a number of forms. Different structures are given
different names e.g. galactose, fructose and glucose, and further differences are shown by a letter
before the molecules name, e.g. α-glucose and β-glucose.

The hydroxyl group, –OH, on carbon atom 1 may be above or below the plane of the ring. The form
of glucose where it is below the ring is known as α-glucose (alpha-glucose) and the form where it is
above the ring is β-glucose (beta-glucose). The same molecule can switch between the two forms.
Two forms of the same chemical are known as isomers, and the extra variety provided by the
existence of α- and β-isomers has important biological consequences, as we shall see in the
structures of starch, glycogen and cellulose.

α-glucose (alpha-glucose) β-glucose (beta-glucose

Disaccharides
Disaccharides, like monosaccharides, are sugars. They are formed by two monosaccharides
joining together. The three most common disaccharides are maltose (glucose + glucose),
sucrose (glucose + fructose) and lactose (glucose + galactose).

The joining of two monosaccharides takes place by a process known as condensation. For
example below two molecules of α-glucose combine to make the disaccharide maltose, or
α-glucose and β-fructose combine to make the disaccharide sucrose. Notice that fructose
has a different ring structure to glucose.

For each condensation reaction, two hydroxyl (–OH) groups line up alongside each other.
One combines with a hydrogen atom from the other to form a water molecule. This allows
an oxygen ‘bridge’ to form between the two molecules, holding them together and forming
a disaccharide (‘di’ means two). The bridge is called a glycosidic bond. In theory any two –
OH groups can line up and, since monosaccharides have many –OH groups, there are a large
number of possible disaccharides. The shape of the enzyme controlling the reaction
determines which –OH groups come alongside each other. Only a few of the possible
disaccharides are common in nature. The reverse of condensation is the addition of water,
which is known as hydrolysis. This takes place during the digestion of disaccharides and
polysaccharides, when they are broken down to monosaccharides.

Roles of Monosaccharides and Disaccharides

Monosaccharides and Disaccharides function as respiratory substrates that are broken down
to provide energy in the form of ATP for carrying out living processes. They are particularly
useful because they have a large number of C—H groups and these can be easily oxidised,
yielding a lot of energy.
Polysaccharides
Polysaccharides are polymers whose monomers are monosaccharides. They are made by
joining many monosaccharides by condensation. Each molecule is joined by a glyosidic bond
with the final containing possible thousands of monosaccharides. The most important ones
are starch, glycogen and cellulose, which are all made of glucose.

Starch
Starch is a polysaccharide found in many parts of a plant in the form of small granules, or
grains, e.g. starch grains in chloroplasts. Starch is a mixture of two substances – amylose and
amylopectin.

• Amylose is composed of 200 to 5000 α-glucose units, which are joined in a straight
chain by 1,4 glycosidic bonds. This chain is then wound into tight coil (like springs)
which makes the molecule more compact and therefore it can be stored more
efficiently as it takes up less space.
• Amylopectin is made up of 5000 to 100000 α-glucose units joined to each other by
1,4 glycosidic bonds which forms shorter chains than amylose and branch out to the
sides. The branches are formed by 1,6 glycosidic bonds.

About 80% of starch is amylopectin and the remaining 20% is amylose. Starch is only
found in plant cells.
Starch is main role is for energy storage, something it is especially suited for because:

• It is insoluble and therefore does not have any osmotic effects within cells, i.e. it
does not tend to draw water into the cells.
• Being insoluble, it does not easily diffuse out of cells.
• It is compact, so a lot of it can be stored in a small space
• When hydrolysed it forms glucose, which is both easily transported and readily
used in respiration, to provide energy in the form of ATP.

Glycogen
Glycogen, like amylopectin, is made of chains of 1,4 linked α-glucose with 1,6 linkages
forming branches. Glycogen molecules tend to be even more branched than
amylopectin molecules. Glycogen molecules clump together to form granules, which are
visible in liver cells and muscle cells, where they form an energy reserve. The smaller
chains of glycogen allow it to hydrolysed quickly to α-glucose. Glycogen is found in
animal cells and fungi but never in plant cells.

Cellulose
Cellulose is the most abundant organic molecule on the planet, due to its
presence in plant cell walls and its slow rate of breakdown in nature. It has a
structural role, being a mechanically strong molecule, unlike starch and
glycogen. However, the only difference between cellulose and starch and
glycogen is that cellulose is a polymer of β-glucose, not α-glucose.
In the β-isomer, the –OH group on carbon atom 1 projects above the ring. In
order to form a glycosidic bond with carbon atom 4, where the –OH group is
below the ring, one glucose molecule must be upside down (rotated 180°)
relative to the other. Thus, successive glucose units are linked at 180° to each
other. This arrangement of β-glucose molecules results in a strong molecule
because the hydrogen atoms of –OH groups are weakly attracted to oxygen
atoms in the same cellulose molecule (the oxygen of the glucose ring) and also to
oxygen atoms of –OH groups in neighbouring molecules. These hydrogen bonds
are individually weak, but so many can form, due to the large number of –OH
groups, that collectively they provide enormous strength. Between 60 and 70
cellulose molecules become tightly cross-linked to form bundles called
microfibrils. Microfibrils are in turn held together in bundles called fibres by
hydrogen bonding.
A cell wall typically has several layers of fibres, running in different directions to
increase strength. Cellulose makes up about 20–40% of the average cell wall;
other molecules help to cross-link the cellulose fibres, and some form a glue-like
matrix around the fibres, which further increases strength.
Cellulose fibres have a very high tensile strength, almost equal to that of steel.
This means that if pulled at both ends they are very difficult to stretch or break
and makes it possible for a cell to withstand the large pressures that develop
within it as a result of osmosis. Without the wall, the cell would burst when in a
dilute solution. These pressures help provide support for the plant by making
tissues rigid and are responsible for cell expansion during growth. The
arrangement of fibres around the cell helps to determine the shape of the cell as
it grows. Despite their strength, cellulose fibres are freely permeable, allowing
water and solutes to reach or leave the cell surface membrane.