All living cells need a supply of oxygen and nutrients to survive.

They also need to remove

waste products so that these do not build up and become toxic. In very small organisms, this
exchange can take place over the surface of the body, so they do not need a specialised
exchange system. In larger organisms with more than two layers of cells, the body surface is
no longer sufficient. Therefore larger organisms need a specialised surface for exchange of
substances with their environment. The three main factors that affect the need for an
exchange system are size, surface area to volume ratio, and level of activity.

Size- In very small organisms (i.e: single-celled organisms), all the cytoplasm is very close to
the environment in which they live, so diffusion supplies enough oxygen and nutrients to
keep the cells alive and active. Multicellular organisms may have several layers of cells. So
any oxygen or nutrients diffusing in from the outside have a longer diffusion pathway.
Diffusion is too slow to enable a sufficient supply to the innermost cells.
Surface area to volume ratio- Small organisms have a small surface area, but they also have
a small volume, so their surface area is large enough to supply all their cells with sufficient
oxygen. As size increases, the volume rises more quickly than the surface area. So large
organisms have a small surface area to volume ratio. Organisms can increase their surface
area by adopting a different shape. An animal such as a flatworm has a very thin, flat body,
giving it a larger surface area to volume ratio (SA:V), but such a body form limits the overall
size that the animal can reach.
Level of activity- Metabolic activity uses energy from food and requires oxygen to release the
energy in aerobic respiration. The cells of an active organism need good supplies of
nutrients and oxygen to supply the energy for movement. This need for energy is increased
in those animals, such as mammals, that keep themselves warm.

All good exchange surfaces have:

A large surface area to provide more space for molecules to pass through. This is often
achieved by folding the walls and membranes involved. A good example is the root hairs in
plants.
A thin barrier to reduce the diffusion distance — and that barrier must be permeable to the
substances being exchanged. This is shown well in the alveoli of the lungs.
A good blood supply to bring fresh supplies of molecules to one side (supply side), keeping
the concentration high, or it may remove molecules from the demand side to keep the
concentration low. This is important to maintain a steep concentration gradient so that
diffusion can occur rapidly. The gills in fish are a good example.

Alveoli: tiny folds of the lung epithelium to increase the surface area.
Bronchi and bronchioles: smaller airways leading into the lungs.
Diaphragm: a layer of muscle beneath the lungs.
Intercostal muscles: muscles between the ribs. Contraction of the external intercostal
muscles raises the ribcage.
Trachea: the main airway leading from the back of the mouth to the lungs.
Ventilation: the refreshing of the air in the lungs, so that there is a higher oxygen
concentration than in the blood, and a lower carbon dioxide concentration.
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concentration than in the blood, and a lower carbon dioxide concentration.

The gaseous exchange system in mammals consists of the lungs and associated airways
that carry air into and out of the lungs. The lungs are a pair of inflatable sacs lying in the
chest cavity. Air can pass into the lungs through the nose and along the trachea (windpipe),
bronchi and bronchioles. Finally, it reaches tiny air-filled sacs called alveoli. These are the
surfaces where the exchange of gases takes place. The lungs are protected by the ribcage.
The ribs are held together by the intercostal muscles. The action of these muscles and the
diaphragm (a layer of muscular tissue beneath the lungs) helps to produce breathing
movements (ventilation).

Gaseous exchange in the lungs

Gases pass by diffusion through the thin walls of the alveoli. Oxygen passes from the air in
the alveoli to the blood in the capillaries. Carbon dioxide passes from the blood to the air in
the alveoli. The lungs must maintain a steep concentration gradient in each direction in order
to ensure that diffusion can continue.

Large surface area to provide more space for molecules to pass through
The individual alveoli are very small — about 100-300 /I,M across. However, they are so
numerous that the total surface area of the lungs is much larger than that of our skin. It has
been calculated that the total surface area of the exchange surface in humans is about 70
m2, or about half the size of a tennis court. The alveoli are lined by a thin layer of moisture,
which evaporates and is lost as we breathe out. The lungs must produce a surfactant that
coats the internal surface of the alveoli to reduce the cohesive forces between the water
molecules, as these forces tend to make the alveoli collapse.

The barrier to exchange is permeable to oxygen and carbon dioxide

The barrier to exchange is comprised of the wall of the alveolus and the wall of the blood
capillary. The cells and their plasma (cell-surface) membranes readily allow the diffusion of
oxygen and carbon dioxide, as the molecules are small and non-polar.

Thin barrier to reduce the diffusion distance

There are a number of adaptations to reduce the distance the gases have to diffuse:
• The alveolus wall is one cell thick.
• The capillary wall is one cell thick.
• Both walls consist of squamous cells — this means flattened or very thin.
• The capillaries are in close contact with the alveolus walls.
• The capillaries are so narrow that the red blood cells are squeezed against the capillary
wall — making them closer to the air in the alveoli and reducing their rate of flow.
So, the total barrier to diffusion is only two flattened cells, and is less than 1µm thick.

A good blood supply

The blood supply helps to maintain a steep concentration gradient, so that the gases
continue to diffuse. • The blood system transports carbon dioxide from the tissues to the
lungs. This ensures that the concentration of carbon dioxide in the blood is higher than that
in the air of the alveoli. Therefore carbon dioxide diffuses into the alveol. The blood also
transports oxygen away from the lungs. This ensures that the concentration of oxygen in the
blood is kept lower than that in the alveoli — so that oxygen diffuses into the blood.

Ventilation The breathing movements ventilate the lungs. This replaces the used air with
fresh air, bringing in more oxygen and removing carbon dioxide. Ventilation ensures that:
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fresh air, bringing in more oxygen and removing carbon dioxide. Ventilation ensures that:
• the concentration of oxygen in the air of the alveolus remains higher than that in the blood
• the concentration of carbon dioxide in the alveoli remains lower than that in the blood.

Therefore, the concentration gradient necessary for diffusion is maintained.

The air we breathe in is not oxygen — it is rich in oxygen. The air we breathe out is not
carbon dioxide — it is rich in carbon dioxide.

Inspiration (inhaling)
the diaphragm contracts to move down and become flatter — this displaces the digestive
organs downwards
the external intercostal muscles contract to raise the ribs
the volume of the chest cavity is increased
the pressure in the chest cavity drops below the atmospheric pressure
air is moved into of the lungs

Expiration (exhaling)
the diaphragm relaxes and is pushed up by the displaced organs underneath
the external intercostal muscles relax and the ribs fall; the internal intercostal muscles can
contract to help push air out more forcefully — this usually only happens during exercise or
coughing and sneezing
the volume of the chest cavity is decreased
the pressure in the lungs increases and rises above the pressure in the surrounding
atmosphere
air is moved out of the lungs

Cartilage: a form of connective tissue. ciliated epithelium: a layer of cells that have many
hair-like extensions called cilia.
Elastic fibres: protein fibres that can deform and then recoil to their original size.
Goblet cells: cells that secrete mucus.
Smooth muscle: involuntary muscle that contracts without the need for conscious thought.

Lung tissue
Alveoli are comprised of squamous epithelium and are surrounded by blood capillaries, so
that the distance that gases must diffuse is very short. The alveolus walls contain elastic
fibres that stretch during inspiration but then recoil to help push air out during expiration. The
alveolus walls are so thin that it may not be possible to distinguish separate cells under a
light microscope.

The airways
To be effective, the airways must meet certain requirements:
• be large enough to allow sufficient air to flow without obstruction
• be supported to prevent collapse when the air pressure inside is low during inspiration
• be flexible in order to allow movement.
The airways are lined by ciliated epithelium, which contributes to keeping the lungs healthy.
Goblet cells in the epithelium release mucus, which traps pathogens. The cilia then move the
mucus up to the top of the airway, where it is swallowed.

The trachea and bronchi

The trachea and bronchus walls have a similar structure. However, the bronchi are narrower
than the trachea. These airways are supported by rings of cartilage which prevent collapse
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than the trachea. These airways are supported by rings of cartilage which prevent collapse
during inspiration. The rings of cartilage in the trachea are C-shaped rather than a complete
ring which allows flexibility and space for food to pass down the oesophagus.

The bronchioles
The bronchioles are much narrower than the bronchi. The larger bronchioles may have
some cartilage, but smaller ones have no cartilage. The wall is comprised mostly of smooth
muscle and elastic fibres. The smallest bronchioles end in clusters of alveoli.
The smooth muscle in the airways does not contribute to the breathing movements.

Smooth muscle and elastic tissue

The smooth muscle can contract. The action of the smooth muscle will constrict the airway.
This makes the lumen of the airway narrower. Constriction of the lumen can restrict the flow
of air to and from the alveoli. Controlling the flow of air to the alveoli might be important if
there are harmful substances in the air. The contraction of the smooth muscle and control of
airflow is not a voluntary act and may occur as a result of an allergic reaction. Once the
smooth muscle has contracted, it cannot reverse this effect on its own. The smooth muscle is
elongated again by the elastic fibres. When the muscle contracts, it deforms the elastic
fibres. As the muscles relax, the elastic fibres recoil to their original size and shape. This acts
to dilate the airway.

Some people overreact to certain substances in the air and their bronchioles constrict
unnecessarily. This is the cause of asthma.

breathing rate: the number of breaths per minute. oxygen uptake: the volume of oxygen
absorbed by the lungs in one minute.
tidal volume: the volume of air inhaled or exhaled in one breath, usually measured at rest.
spirometer: a device that can measure the movement of air into and out of the lungs.
vital capacity: the greatest volume of air that can be expelled from the lungs after taking the
deepest possible breath.

Using a spirometer
Lung volumes can be measured using a spirometer. This is a device that measures the
movement of air in and out of the lungs as a person breathes. A float-chamber spirometer
consists of a chamber of air or medical-grade oxygen floating on a tank of water. During
inspiration, air is drawn from the chamber so that the lid moves down. During expiration, the
air returns to the chamber, raising the lid. These movements may be recorded on a data
logger.
The carbon dioxide-rich air exhaled is passed through a chamber of soda lime, which
absorbs the carbon dioxide. This allows the measurement of oxygen consumption.
Precautions that must be taken when using a spirometer:
• The subject should be healthy and, in particular, free from asthma
• The soda lime should be fresh and functioning.
• There should be no air leaks in the apparatus, as this would give invalid or inaccurate
results.
• The mouthpiece should be sterilised.
• The water chamber must not be overfilled (or water may enter the air tubes).

Modern spirometers may be small and simple hand-held devices, These record the
movements of air in and out of the lungs. However, many cannot measure the rate of oxygen
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movements of air in and out of the lungs. However, many cannot measure the rate of oxygen
consumption.

Lung volumes
The total lung volume consists of the vital capacity, which can be measured, and the residual
volume, which cannot be measured using the spirometer.
• Vital capacity is the maximum volume of air that can be moved by the lungs in one breath.
This is measured by taking a deep breath and expiring all the air possible from the lungs.
Vital capacity depends upon a number of factors such as:
the size of a person (particularly their height)
their age and gender
their level of regular exercise.
Vital capacity is usually in the region of 2.5-5.0 dm3, but may rise above this in trained
athletes.

• The residual volume is the volume of air that remains in the lungs even after forced
expiration. This air remains in the airways and alveoli. This is approximately 1.5 dm3.
• Tidal volume is the volume of air moved in and out with each breath. It is normally
measured at rest. A typical tidal volume at rest might be 0.5 dm3. This is usually sufficient to
supply all the oxygen required in the body at rest.

Oxygen uptake
As carbon dioxide is removed using soda lime, the volume of air in the chamber decreases.
This decrease can be observed and measured on the spirometer trace. We can assume that
the volume of carbon dioxide released and absorbed by the soda lime equals the volume of
oxygen absorbed by the blood. Therefore, measuring the gradient of the decrease in volume
enables us to calculate the rate of oxygen uptake.

Calculating oxygen uptake from a spirometer trace

• On trace, draw a line from the initial oxygen volume (A) down to the horizontal axis, and
another line from the final oxygen volume (B) to the horizontal axis. Measure the length of
time between these points
• Measure the difference in volume between points A and B
• Divide by the time taken for this decrease
• The unit will be dm3s-1.

Breathing rate and oxygen uptake

The breathing rate can also be measured from a spirometer trace. Simply count the number
of peaks in each minute. Breathing rate at rest is usually about 12-14 breaths per minute.
Oxygen uptake will depend upon a number of factors. A higher oxygen uptake will result from
increased demand, such as during exercise when the muscles are respiring more.
Increased oxygen uptake will result from:
• increased breathing rate
• deeper breaths

Bony fish
Bony fish must exchange gases with the water in which they live. They use gills in order to
absorb oxygen dissolved in the water and release carbon dioxide into the water. The oxygen
concentration will be typically much lower than is found in air. Most bony fish have five pairs
of gills (see Figure 1) which are covered by a bony plate called the operculum. Each gill
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consists of two rows of gill filaments (primary lamellae) attached to a bony arch. The
filaments are very thin, and their surface is folded into many secondary lamellae (or gill
plates). This provides a very large surface area. Blood capillaries carry deoxygenated blood
close to the surface of the secondary lamellae where exchange takes place.

Countercurrent flow
Blood flows along the gill arch and out along the filaments to the secondary lamellae. The
blood then flows through capillaries in the opposite direction to the flow of water over the
lamellae. This arrangement creates a countercurrent flow that absorbs the maximum amount
of oxygen from the water

Ventilation in bony fish

Bony fish can keep water flowing over the gills by using a buccal—opercular pump. The
buccal cavity (mouth) can change volume. The floor of the mouth moves downwards,
drawing water into the buccal cavity. The mouth closes and the floor is raised again pushing
water through the gills. Movements of the operculum are coordinated with the movements of
the buccal cavity As water is pushed from the buccal cavity, the operculum moves outwards.
This movement reduces the pressure in the opercular cavity (the space under the
operculum), helping water to flow through the gills.

Insects
Insects do not transport oxygen in blood. Insects have an open circulatory system in which
the body fluid acts as both blood and tissue fluid. Circulation is slow and can be affected by
body movements.
Insects possess
3.1 an air-filled tracheal system, which supplies air directly to all the respiring
tissues. Air enters the system via a pore in each segment, called a spiracle. The air is
transported into the body through a series of tubes called tracheae (singular `trachea').
These divide into smaller and smaller tubes, called tracheoles. The ends of the tracheoles
are open and filled with fluid called tracheal fluid. Gaseous exchange occurs between the air
in the tracheole and the tracheal fluid. Some exchange can also occur across the thin walls
of the tracheoles.

Many insects are very active and need a good supply of oxygen. When tissues are active,
the tracheal fluid can be withdrawn into the body in order to increase the surface area of the
tracheole wall exposed to air. This means that more oxygen can be absorbed when the
insect is active.

Ventilation in insects
Larger insects can also ventilate their tracheal system by movements of the body. This can
be achieved in a number of ways.
• In many insects, sections of the tracheal system are expanded and have flexible walls.
These act as air sacs which can be squeezed by the action of the flight muscles. Repetitive
expansion and contraction of these sacs ventilate the tracheal system.
• In some insects, movements of the wings alter the volume of the thorax. As the thorax
volume decreases, air in the tracheal system is put under pressure and is pushed out of the
tracheal system. When the thorax increases in volume, the pressure inside drops and air is
pushed into the tracheal system from outside.
• Some insects have developed this ventilation even further. Locusts can alter the volume of
their abdomen by specialised breathing movements. These are coordinated with opening
and closing valves in the spiracles. As the abdomen expands, spiracles at the front end of
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and closing valves in the spiracles. As the abdomen expands, spiracles at the front end of
the body open and air enters the tracheal system. As the abdomen reduces in volume, the
spiracles at the rear end of the body open and air can leave the tracheal system.
The mechanism of ventilation in insects is similar to that in mammals. An increase in the
volume of the body cavity reduces pressure so that air enters the tracheal system; a
decrease in volume raises the pressure to push air out again.
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