Chapter 8

Transport in
mammals

LEARNING INTENTIONS
In this chapter you will learn how to:
• describe the structure of the mammalian circulatory system
• explain how the structures of arteries, arterioles, veins, venules and capillaries are related to their functions
• describe and explain the structure and functions of blood, including the transport of oxygen and carbon
dioxide
• make diagrams of blood vessels and blood cells from slides, photomicrographs or electron micrographs
• describe the formation and functions of tissue fluid
• explain the structure and function of the heart
• describe the cardiac cycle and its control.

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 8 Transport in mammals

BEFORE YOU START

• In your group, make lists of:
– the different kinds of blood vessel in the mammalian circulatory system
– the different components of blood.
• Write two facts about each of the items in your lists – for example, what their function is, and how they
are adapted to carry it out.
Be ready to share your ideas with others in the class.

ARTIFICIAL HEARTS
Each year, about 18 million people worldwide die
from cardiovascular disease, more than from any
other disease. ‘Cardiovascular’ means to do with the
heart and the circulatory system, and many of these
deaths are due to the heart failing to work normally.
In many countries, medical help is available for
people with a failing heart, ranging from treatment
with drugs to major heart surgery. But, until recently,
the only hope for some heart patients was a heart
transplant. However, the number of people needing
a new heart is much greater than the number of
hearts available. Many people wait years for a heart
transplant, and many die from their heart disease
before they get a new heart.
Petar Bilic (not his real name) thought that he was
going to add to that statistic. The muscle in both
of his ventricles had deteriorated so much that his
heart was only just keeping him alive. No suitable Figure 8.1: An artificial heart.
heart could be found for a transplant. Biomedical engineers continue to make progress
Petar was very lucky. In recent years, biomedical in developing new types of heart that should work
engineers have developed a pumping device called for much longer – perhaps long enough for its
a ‘total artificial heart’ (Figure 8.1). Petar’s heart was owner to live out a long life without any need for a
completely removed, and an artificial heart put in heart transplant. And, while the first artificial hearts
its place. Petar was able to go home within a few were designed to fit into the body of an adult man,
weeks of his operation. The plan is that the artificial smaller ones are now available, which can be used
heart will keep him alive until a real heart is available for women and children.
for transplant. Some patients have lived for almost Question for discussion
5 years with their artificial heart in place. However,
living with an artificial heart is not easy. An artificial What do you think might be the advantages and
heart needs an energy supply, and this is often disadvantages of using an artificial heart rather than
provided with a battery that the patient carries in a a heart transplant, to treat a person whose own
backpack. heart is failing?

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Key lungs
8.1 Transport systems oxygenated blood
deoxygenated blood
in animals RA right atrium
pulmonary
circulation
LA left atrium
Most animals are far more active than plants. They must
move to find their food, because they cannot make their RV right ventricle
RA LA
own. Movement requires energy, for example for muscle LV left ventricle
RV LV
contraction and the transmission of nerve impulses. This
energy comes from glucose and other substances, which systemic
are broken down in respiration inside each individual cell. circulation

The most efficient form of respiration – which releases

the most energy from a given amount of glucose – is rest of
aerobic respiration, and this requires good supplies of
right side of body body left side of body
oxygen. Supplying oxygen to respiring tissues is one of
the most important functions of an animal’s transport
system. At the same time, waste products such as carbon Figure 8.2: The general plan of the mammalian transport
dioxide can be removed. system, viewed as though looking at someone facing you. It
is a closed double circulatory system.
Very small animals may be able to get enough oxygen
to their cells by diffusion, especially if they are not
Blood is pumped out of the left ventricle into the aorta
particularly active. In a jellyfish, for example, oxygen
(Figure 8.3), and travels from there to all parts of the
simply diffuses into its body from the seawater around
body except the lungs. It returns to the right side of
it, and then to the respiring cells. Carbon dioxide
the heart in the vena cava. This is called the systemic
diffuses in the opposite direction. Because no cell is very
circulation.
far from the surface, each cell gets an adequate amount
of oxygen quickly enough for its needs. The blood is then pumped out of the right ventricle into
the pulmonary arteries, which carry it to the lungs. The
But in larger animals, such as mammals, diffusion is not
final part of the journey is along the pulmonary veins,
sufficient. A transport system is needed to distribute
which return it to the left side of the heart. This is called
oxygen quickly to all the body cells, and to remove their
the pulmonary circulation.
waste products. Mammals have greater requirements
for oxygen than most other animals because they use
respiration to generate heat inside their bodies, to help KEY WORDS
to keep their body temperature constant.
circulatory system: a system that carries fluids
around an organism’s body

8.2 The mammalian closed blood system: a circulatory system made

up of vessels containing blood

circulatory system double circulation: a circulatory system in which

the blood passes through the heart twice on one
Figure 8.2 shows the general layout of the main complete circuit of the body
transport system of mammals – the blood system or
circulatory system. It is made up of a pump – the heart – systemic circulation: the part of the circulatory
and a system of interconnecting tubes – the blood system that carries blood from the heart to all of
vessels. The blood always remains within these vessels, the body except the gas exchange surface, and
and so the system is known as a closed blood system. then back to the heart
Put your finger onto the left ventricle in Figure 8.2. Use pulmonary circulation: the part of the circulatory
your finger to follow the journey of the blood around system that carries blood from the heart to
the body. You will find that the blood travels twice the gas exchange surface and then back to
through the heart on one complete ‘circuit’. This is the heart
called a double circulation.

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 8 Transport in mammals

The pressure in the systemic circulation is considerably gills

higher than in the pulmonary circulation. You can read
about blood pressure in the circulatory system later in
this chapter.
ventricle
jugular vein carotid artery
subclavian vein subclavian artery
atrium
pulmonary aorta
vein

rest of
heart
body

pulmonary artery Figure 8.4: The general plan of the transport system of
vena cava
a fish.
hepatic vein lung
liver
hepatic artery
hepatic
portal vein kidney 8.3 Blood vessels
gut
renal artery There are three main types of vessel making up the
renal vein circulatory system. Figure 8.5 shows these vessels in
mesenteric artery
iliac vein transverse section. Vessels carrying blood away from the
iliac artery
heart are known as arteries, while those carrying blood
towards the heart are veins. Small arteries are called
arterioles, and small veins are venules. Linking arterioles
Figure 8.3: The positions of some of the main blood and venules, taking blood close to almost every cell in
vessels in the human body. the body, are tiny vessels called capillaries.

KEY WORDS
Question
artery: vessel with thick, strong walls that carries
1 Figure 8.4 shows the general layout of the
high-pressure blood away from the heart
circulatory system of a fish.
With a partner, discuss: vein: vessel with relatively thin walls that carries
low-pressure blood back to the heart
a how this system differs from the circulatory
system of a mammal arteriole: small artery
b why the mammalian transport system may be venule: small vein
able to deliver more oxygen more quickly to capillary: the smallest blood vessel, whose role is
the tissues than the fish’s transport system to deliver oxygen and nutrients to body tissues,
c how these differences could relate to the and to remove their waste products
different requirements of a fish and a mammal.
Be ready to share your ideas with the rest of the class.

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Transverse section (TS) through small artery

inner layer, which is the endothelium Arteries in different parts of the body
(a very smooth, single layer of cells – vary in their structure. Arteries near
squamous epithelium) the heart have especially large
numbers of elastic fibres in the middle
relatively narrow lumen layer, as shown here. In other parts of
the body, the middle layer contains
middle layer, containing elastic fibres, less elastic tissue and more smooth
collagen fibres and smooth muscle muscle.

outer layer, containing collagen

fibres and some elastic fibres
TS through capillary
7 μm

TS through small vein wall made of

endothelium,
one cell thick
lumen, just big enough
for a red cell to squeeze
inner later, like that along
of the artery

relatively large lumen

middle layer, very thin,

containing some smooth
0.7 mm muscle and elastic fibres

outer layer, mostly

collagen fibres

Figure 8.5: The tissues making up the walls of arteries, capillaries and veins.

Arteries and arterioles KEY WORDS

The function of arteries is to transport blood, swiftly and
endothelium: a tissue that lines the inner surface
at high pressure, to the tissues.
of a structure such as a blood vessel
Artery walls are very strong and elastic. Blood leaving squamous epithelium: one or more layers of
the heart is at a very high pressure. Blood pressure in the thin, flat cells forming the lining of some hollow
human aorta may be around 120 mmHg, which can also structures, e.g. blood vessels and alveoli
be stated as 16 kPa. The thickness and composition of
the artery wall enables it to withstand this pressure. smooth muscle: a type of muscle that can
contract steadily over long periods of time
Both arteries and veins have walls made up of three
layers (Figures 8.5 and 8.6):
• an inner layer, which is made up of a layer of IMPORTANT
endothelium (lining tissue) consisting of a layer
of flat cells (squamous epithelium) fitting together Blood pressure is still measured in the old units
like jigsaw pieces, plus a layer of elastic fibres; the of mmHg, even though kPa is the SI unit. The
endothelium is very smooth, minimising friction abbreviation mmHg stands for ‘millimetres of
with the moving blood mercury’, and refers to the distance which a
column of mercury is pushed up the arm of a
• a middle layer containing smooth muscle, collagen U-tube. 1 mmHg is equivalent to about 0.13 kPa.
and elastic fibres
• an outer layer containing elastic fibres and collagen
fibres.

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 8 Transport in mammals

Figure 8.6: Photomicrograph of an artery (left) and a vein (right) (×110).

Arteries have the thickest walls of any blood vessel. are not entirely effective in achieving this: if you feel
The aorta, the largest artery, has an overall diameter your pulse in your wrist, you can feel the artery, even at
of 2.5 cm close to the heart, and a wall thickness of this distance from your heart, being stretched outwards
about 2 mm. The composition of the wall provides great with each surge of blood from the heart.
strength and resilience. The middle layer, which is by far
As arteries divide into smaller vessels, the proportion
the thickest part of the wall, contains a large amount of
of muscle in their walls increases and the proportion
elastic fibres. These allow the wall to stretch as pulses of
of elastic tissue decreases. They are now muscular
blood surge through at high pressure. Arteries further
arteries. Muscular arteries take blood from an elastic
away from the heart have fewer elastic fibres in the
artery and deliver it close to its final destination. The
middle layer but have more muscle fibres.
type of muscle in their walls is smooth muscle, which is
Arteries that have a lot of elastic tissue in their middle able to contract slowly and steadily to alter the internal
layer – such as the aorta – are called elastic arteries. diameter of the artery and therefore control the volume
The function of an elastic artery is to carry blood from of blood that can flow through it.
the heart on the first part of its journey towards its
Muscular arteries divide to form even smaller vessels
final destination. The elasticity of these artery walls is
called arterioles. These also contain a lot of smooth
important in allowing them to stretch, which reduces
muscle in their walls. Their narrowness provides
the likelihood that they will burst. This elasticity also
has another very important function. Blood is pumped
out of the heart in pulses, rushing out at high pressure
KEY WORDS
as the ventricles contract, and slowing as the ventricles elastic arteries: relatively large arteries, which
relax. The artery walls stretch as the high-pressure blood have a lot of elastic tissue and little muscle tissue
surges into them, and then recoil inwards as the pressure in their walls
drops. Therefore, as blood at high pressure enters an
artery, the artery becomes wider, reducing the pressure muscular arteries: arteries that are closer to the
a little. As blood at lower pressure enters an artery, the final destination of the blood inside them than
artery wall recoils inwards, giving the blood a small elastic arteries, with more smooth muscle in their
‘push’ and raising the pressure a little. The overall effect walls which allows them to constrict and dilate
is to ‘even out’ the flow of blood. However, the arteries

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platelet red blood cell throughout every tissue in the body except the brain,
inner layer, cornea and cartilage. Such networks are sometimes
containing called capillary beds.
endothelial
cells The small size of capillaries is of great importance in
allowing them to bring blood as close as possible to
elasc fibres in each group of cells in the body. A human capillary is
the inner approximately 7 μm in diameter, about the same size as
layer
a red blood cell (Figures 8.5, 8.8 and 8.9). The walls of
middle layer, capillaries are extremely thin because they are made up
containing of a single layer of endothelial cells. As red blood cells
smooth carrying oxygen squeeze through a capillary, they are
muscle ssue brought to within as little as 1 μm of the cells outside the
capillary that need the oxygen.
outer layer,
containing
collagen

Figure 8.7: Transmission electron micrograph (TEM) of a

small artery.

resistance to blood flow, causing it to slow down, which

provides extra time for exchange of gases and nutrients
as the blood flows through the capillaries in the tissues.
The walls of arterioles have a nerve supply. Nerve
impulses from the brain can cause their smooth muscle
to contract, narrowing the arteriole. This is called
vasoconstriction. This can be used to reduce blood flow
to a particular area and divert it to other tissues. When
the muscle relaxes, the diameter of the arteriole widens.
red blood cell endothelial cell
This is called vasodilation. The smooth muscle can also
respond to hormones in the blood. Figure 8.8: Photomicrograph of a blood capillary
containing red blood cells (dark red) (×900). The cells of
KEY WORDS the endothelium are very thin, except where there is a
nucleus (red).
vasoconstriction: the narrowing of a muscular
artery or arteriole, caused by the contraction of In most capillaries, there are tiny gaps between the
the smooth muscle in its walls individual cells that form the endothelium. As you will see
vasodilation: the widening of a muscular artery later in this chapter, these gaps are important in allowing
or arteriole, caused by the relaxation of the some components of the blood to seep through into the
smooth muscle in its walls spaces between the cells in all the tissues of the body.
By the time blood reaches the capillaries, it has already
lost much of the pressure originally supplied to it by the
contraction of the ventricles. Blood pressure continues
Capillaries to drop as it passes through the capillaries. As blood
enters a capillary from an arteriole, it may have a
The arterioles themselves continue to branch, eventually pressure of around 35 mmHg or 4.7 kPa; by the time it
forming the tiniest of all blood vessels, capillaries. reaches the far end of the capillary, the pressure will have
The function of capillaries is to take blood as close as dropped to around 10 mmHg or 1.3 kPa.
possible to all cells, allowing rapid transfer of substances
between cells and blood. Capillaries form a network

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 8 Transport in mammals

Question several leg muscles. Whenever you tense (contract) these

muscles, they squeeze inwards on the veins in your legs,
2 Suggest why there are no blood capillaries in temporarily raising the pressure within them.
the cornea of the eye. How might the cornea be This squeezing, in itself, would not help to push the
supplied with oxygen and nutrients? blood back towards the heart; blood would just squidge
capillary wall, up and down as you walked. To keep the blood flowing
nucleus of made of a single layer in the right direction, veins contain half-moon valves,
endothelial cell of endothelial cells or semilunar valves, formed from their endothelium
(Figure 8.10). These valves allow blood to move
towards the heart, but not away from it. So, when
you contract your leg muscles, the blood in the veins
is squeezed up through these valves, but cannot pass
down through them.

to heart

Pressure in
the vein is semilunar valve,
produced preventing flow
by skeletal of blood away
muscles from heart
contracting
mitochondrion red blood cell close to it.
in endothelial in the lumen
cell of the capillary
Figure 8.10: Longitudinal section (LS) through part of a
Figure 8.9: TEM of a transverse section (TS) through a small
small vein.
capillary (approximately ×4500).

Veins and venules KEY WORD

As blood leaves a capillary bed, the capillaries gradually semilunar valve: a half-moon shaped valve,
join with one another, forming larger vessels called such as the ones in the veins and between the
venules. These join to form veins. The function of veins ventricles and arteries
is to return blood to the heart.
By the time blood enters a vein, its pressure has dropped
to a very low value. In humans, a typical value for
venous blood pressure is about 5 mmHg or less. This Blood pressure in the
very low pressure means that there is no need for veins
to have thick walls. They have the same three layers as circulatory system
arteries, but the middle layer is much thinner and has You have seen how blood leaves the heart at high
far fewer elastic fibres and muscle fibres. pressure, and then gradually loses this pressure as it
The low blood pressure in veins creates a problem: passes through muscular arteries, arterioles, capillaries,
how can this blood be returned to the heart? Think venules and veins. This happens in both systems –
about how blood can return to your heart from your the systemic system and the pulmonary system. The
feet when you are standing up. Unaided, the blood in pressure of blood leaving the heart is much greater in
your leg veins would sink and accumulate in your feet.
However, many of the veins run within, or very close to,

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120

100
Blood pressure / mmHg

80

60

40

20

0
aorta arteries arterioles capillaries venules veins venae pulmonary arterioles capillaries venules pulmonary
cavae arteries veins

Systemic circulation Pulmonary circulation

Figure 8.11: Blood pressure in different regions of the human circulatory system.

the systemic system than in the pulmonary system. • Who has written the best reasons? What
These blood pressure changes are shown in Figure 8.11. makes these the best?
If you think your table can be improved, make
changes to it.
Questions
3 Suggest reasons for each of the following.
a Normal venous pressure in the feet is about
25 mmHg. When a soldier stands motionless
8.4 Tissue fluid
at attention, the blood pressure in his feet rises Blood is composed of cells floating in a pale yellow
very quickly to about 90 mmHg. liquid called plasma. Blood plasma is mostly water, with
a variety of substances dissolved in it. These solutes
b When you breathe in – that is, when the
include nutrients such as glucose and waste products
volume of the thorax increases – blood moves
such as urea that are being transported from one place
through the veins towards the heart.
to another in the body. Solutes also include protein
4 Using the graph in Figure 8.11, describe and molecules, called plasma proteins, which remain in the
explain in your own words how blood pressure blood all the time.
varies in different parts of the circulatory system.
5 a Construct a table comparing the structure of
KEY WORDS
arteries, veins and capillaries. Include both
similarities and differences, and give reasons plasma: the liquid component of blood, in which
for the differences you describe. the blood cells float; it carries a very large range
b Compare your table with others. of different substances in solution
• Are the headings of the rows and columns plasma proteins: a range of several different
the same? If not, whose do you think are proteins dissolved in the blood plasma, each with
best, and why? their own function; many of them are made in
• Has anyone else thought of a similarity or the liver
difference that you did not?

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 8 Transport in mammals

As blood flows through capillaries within tissues, some water moves from the capillaries into the tissue fluid
of the plasma leaks out through the gaps between the (Figure 8.12).
cells in the walls of the capillary, and flows gently into
At the venule end of a capillary bed, the blood pressure
the spaces between the cells of the tissues. Almost
inside the capillaries is lower, so there is less tendency
one-sixth of your body consists of spaces between your
for water to be pushed out of the capillaries into the
cells. These spaces are filled with this leaked plasma,
tissue. The water potential gradient caused by the
which is known as tissue fluid.
difference in the concentration of dissolved proteins
is still similar to that at the arteriole end. Now, the net
KEY WORD movement of water is from the tissue fluid, back into
tissue fluid: the almost colourless fluid that fills the capillaries.
the spaces between body cells; it forms from the Overall, more fluid flows out of capillaries than into
fluid that leaks from blood capillaries them, so there is a net loss of fluid from the blood as it
flows through a capillary bed.
Tissue fluid is almost identical in composition to blood If blood pressure is too high, too much fluid is forced
plasma. However, it contains far fewer protein molecules out of the capillaries and may accumulate in the tissues.
than blood plasma, because these are too large to escape This build-up of fluid is called oedema. One of the roles
easily through the capillary endothelium. Red blood of arterioles is to reduce the pressure of the blood that
cells are much too large to pass through, so tissue fluid enters the capillaries, in order to avoid this.
does not contain these, but some white blood cells can
Tissue fluid forms the environment of each individual
squeeze through and move around freely in tissue fluid.
body cell. Exchanges of materials between cells and
The volume of fluid that leaves the capillary to form the blood occur through the tissue fluid. Within
tissue fluid is the result of two opposing forces. At the your body, many processes take place to maintain
arterial end of a capillary bed, the blood pressure inside the composition of tissue fluid at a constant level, to
the capillary is enough to push fluid out into the tissue. provide an optimum environment in which cells can
However, there is a greater concentration of dissolved work. These processes contribute to the overall process
proteins in the blood plasma than in the tissue fluid. of homeostasis – that is, the maintenance of a constant
This produces a water potential gradient from the tissue internal environment – and include the regulation of
fluid into the blood plasma (see Chapter 4, Section 4.5, glucose concentration, water, pH, metabolic wastes and
Movement of substances across membranes). Overall, temperature.

cells

arterial end of capillary blood venous end of capillary

plasma

tissue fluid

solute concentration hydrostatic pressure solute concentration hydrostatic pressure

gradient gradient gradient gradient

net movement of water out net movement of water in

Figure 8.12: Movement of fluid into and out of capillaries.

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Questions intestine to the liver, and from the liver to all other body
cells. Urea is transported from the liver to the kidneys.
6 Table 8.1 shows the relative permeability of Water’s properties as a solvent (Chapter 2, Section 2.7,
capillary walls in a muscle to different substances. Water) make it ideal for this role.
In the table, the permeability of water is given As well as substances in solution, blood plasma
a value of 1, and the other values are given in transports heat around the body. You will remember
proportion to this. that water has a high heat capacity, which allows it
Use the information in the table, and your to absorb a lot of heat energy without altering its
own knowledge, to discuss whether there is a temperature very much.
relationship between the RMM of a substance As tissue fluid is formed from blood plasma, it also
and the permeability of the capillary walls to contains a high percentage of water. The high heat
that substance. If so, how can you explain this capacity of the water in tissue fluid helps the whole body
relationship? to maintain a relatively constant temperature.
Substance Relative molecular Permeability of
mass (RMM) capillary walls a

water 18 1.00
sodium ions 23 0.96
urea 60 0.8
glucose 180 0.6
haemoglobin 68 000 0.01
albumin 69 000 0.000 01

Table 8.1: Relative permeability of capillary walls in muscle.

7 The most abundant plasma protein is albumin.

Suggest why it is important that capillary walls are b red blood cell lymphocyte platelets
not permeable to albumin.
8 The disease kwashiorkor is caused by a diet which
is very low in protein. The concentration of
proteins in blood plasma is much lower than usual.
One of the symptoms of kwashiorkor is swelling
caused by build-up of tissue fluid. Suggest why this
is so.

8.5 Blood
monocyte (macro- neutrophils – one kind of
You have about 5 dm3 of blood in your body, with a phage) – large endocytic white blood cell,
mass of about 5 kg. Suspended in the blood plasma, endocytic (phagocytic) containing a nucleus with two
you have around 2.5 × 1013 red blood cells, 5 × 1011 white cell with a large oval or to eight lobes and many pink
kidney-shaped nucleus granules in the cytoplasm
blood cells and 6 × 1012 platelets (small cell fragments
with no nucleus) (Figures 8.13 and 8.14).
Figure 8.13: a Photomicrograph of human blood. It has
Blood plasma is a pale yellow liquid. It is about 95% been stained so that the nuclei of the cells are dark purple
water, with various substances dissolved in it. These (×1600). b Diagram of the types of cells seen in a stained
substances are transported in the blood from one blood film.
part of the body to another. For example, glucose is
transported, in solution in blood plasma, from the small

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 8 Transport in mammals

Red blood cells The structure of a red blood cell is unusual in several ways.
• Red blood cells are shaped like a biconcave disc.
The red colour of red blood cells (Figures 8.14 and
The dent in each side of the cell increases the
8.15) is caused by the pigment haemoglobin, a globular
surface area to volume ratio (surface area : volume)
protein (Chapter 2, Section 2.6, Proteins). The main
of the cell. This large surface area means that
function of haemoglobin is to transport oxygen from
oxygen can diffuse quickly into or out of the cell.
lungs to respiring tissues. This function is described in
detail later in this chapter. • Red blood cells are very small. The diameter of a
human red blood cell is about 7 μm, compared with
the diameter of an average liver cell of 40 μm. This
small size means that no haemoglobin molecule
within the cell is very far from the cell surface
membrane, and the haemoglobin molecules can
therefore quickly exchange oxygen with the fluid
outside the cell. It also means that capillaries can
be only 7 μm wide and still allow red blood cells to
squeeze through them, so bringing oxygen as close
as possible to cells which require it.
• Red blood cells are very flexible. Some capillaries
are even narrower than the diameter of a red blood
cell. The cells are able to be squashed so that they
can pass through these vessels. This is possible
because the cells have a specialised cytoskeleton
(Chapter 1, Section 1.6, Plant and animal cells as
seen with an electron microscope), made up of a
mesh-like network of protein fibres. This allows
Figure 8.14: False-colour scanning electron micrograph of
them to be squashed into different shapes but then
human blood. Red blood cells have been coloured red. The
spring back to produce the normal biconcave shape.
blue spheres are white blood cells. The platelet has been
coloured yellow. • Red blood cells have no nucleus, no mitochondria
and no endoplasmic reticulum. The lack of these
organelles means that there is more room for
haemoglobin, so maximising the amount of oxygen
cell surface membrane
which can be carried by each red blood cell.
Red blood cells do not live very long. Old ones are
haemoglobin solution in TS through a red blood cell
broken down in the liver, and new ones are constantly
cytoplasm (no nucleus, made in the bone marrow.
mitochondria or ER)

Questions
9 Assuming that you have 2.5 × 1013 red blood cells in
your body, that the average life of a red blood cell
is 120 days, and that the total number of red blood
cells remains constant, calculate how many new red
7 μm blood cells must be made, on average, in your bone
marrow each day.
10 Which of these functions could, or could not, be
Figure 8.15: Red blood cells. carried out by a red blood cell? In each case, briefly
justify your answer.
a protein synthesis c lipid synthesis
b cell division d active transport

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White blood cells antibodies, which attach to and destroy the invading
cells. There are different types of lymphocyte, which act
White blood cells, like red blood cells, are made in the in different ways, although they all look the same. Their
bone marrow. They are easy to distinguish from red activities are described in Chapter 11 (Section 11.2, Cells
blood cells in a micrograph because: of the immune system). Lymphocytes are smaller than
most phagocytes, and they have a large round nucleus
• white blood cells all have a nucleus, although the
and only a small amount of cytoplasm.
shape of this varies in different types of white cell
• most white blood cells are larger than red blood
cells, although one type, lymphocytes, may be KEY WORDS
slightly smaller
neutrophil: one type of phagocytic white blood
• white blood cells are either spherical or irregular in cell; it has a lobed nucleus and granular cytoplasm
shape, not a biconcave disc (compare Figures 8.14 monocyte: the largest type of white blood cell; it
and 8.16). has a bean-shaped nucleus; monocytes can leave
There are many different kinds of white blood cell, with the blood and develop into a type of phagocytic
a wide variety of functions, although all are concerned cell called a macrophage
with fighting disease. They can be divided into two main macrophage: phagocytic cell found in tissues
groups: phagocytes and lymphocytes. throughout the body; they act as antigen-
Phagocytes are cells that destroy invading presenting cells (APCs)
microorganisms by phagocytosis. The commonest lymphocyte: a white blood cell with a nucleus
type of phagocyte is called a neutrophil, and can be that almost fills the cell, which responds to
recognised by its lobed nucleus and granular cytoplasm. antigens and helps to destroy the antigens or the
Monocytes (Figure 8.13) are cells that can develop into a structure that is carrying them
different type of phagocyte called a macrophage.

PRACTICAL ACTIVITY 8.1

Observing and drawing blood vessels and

blood cells
You should practise using prepared microscope
slides to identify sections of arteries and veins.
Plan diagrams can be used to show the different
tissue layers in the walls of blood vessels. Look
back at Practical Activity 7.1 in Chapter 7 to
remind yourself how to make plan diagrams.
You can also use prepared slides, electron
micrographs and photomicrographs to observe
and draw blood cells – red cells, monocytes,
neutrophils and lymphocytes. If you are viewing a
prepared microscope slide, you will need to use
high power, and your drawings will be high-power
details, showing the structures of individual cells.
Figure 8.16: False-colour scanning electron micrograph Look back at Practical Activity 7.1 for advice on
of a section through a white blood cell (×6000). This is a high-power detail drawings, and some examples.
lymphocyte.
(See Practical Investigations 7.1 and 7.3 in the
Lymphocytes also destroy microorganisms, but not by Practical Workbook for additional information.)
phagocytosis. Some of them secrete chemicals called

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 8 Transport in mammals

Haemoglobin To investigate how haemoglobin behaves, samples

are extracted from blood and exposed to different
A major role of the circulatory system is to transport concentrations, or partial pressures, of oxygen. The
oxygen from the gas exchange surfaces of the alveoli in quantity of oxygen which combines with each sample of
the lungs to tissues all over the body. Body cells need a haemoglobin is then measured. The maximum amount
constant supply of oxygen in order to be able to carry of oxygen with which a sample can possibly combine
out aerobic respiration. Oxygen is transported around is given a value of 100%. A sample of haemoglobin
the body inside red blood cells in combination with the which has combined with this maximum amount of
protein haemoglobin (Figure 2.23). oxygen is said to be saturated. The amounts of oxygen
As you saw in Chapter 2 (Section 2.6, Proteins), each with which identical samples of haemoglobin combine
haemoglobin molecule is made up of four polypeptides, at lower oxygen partial pressures are then expressed
each containing one haem group. Each haem group can as a percentage of this maximum value known as the
combine with one oxygen molecule, O2. Overall, then, percentage saturation. Table 8.2 shows a series of results
each haemoglobin molecule can combine with four from such an investigation.
oxygen molecules (eight oxygen atoms). The percentage saturation of each sample can be
4O2 HbO8 plotted against the partial pressure of oxygen to obtain
Hb + →
the curve shown in Figure 8.17. This is known as a
haemoglobin oxygen oxyhaemoglobin dissociation curve.
The curve shows that at low partial pressures of oxygen,
Question the percentage saturation of haemoglobin is very low –
that is, the haemoglobin is combined with only a very
11 In a healthy adult human, there is about 150 g of little oxygen. At high partial pressures of oxygen, the
haemoglobin in 1 dm3 of blood. percentage saturation of haemoglobin is very high; it is
a 1 g of pure haemoglobin can combine with combined with large amounts of oxygen.
1.3 cm3 of oxygen at body temperature.
Calculate how much oxygen can be carried in
1 dm3 of blood. KEY WORDS
b At body temperature, the solubility of oxygen partial pressure: a measure of the concentration
in water is approximately 0.025 cm3 of oxygen of a gas
per cm3 of water. Assuming that blood plasma
is mostly water, how much oxygen could be percentage saturation: the degree to which
carried in 1 dm3 of blood if there was no the haemoglobin in the blood is combined
haemoglobin? with oxygen, calculated as a percentage of the
maximum amount with which it can combine
The haemoglobin dissociation curve dissociation curve: a graph showing the
A molecule whose function is to transport oxygen from percentage saturation of a pigment (such as
one part of the body to another must be able to pick up haemoglobin) with oxygen, plotted against the
oxygen efficiently at the lungs. It is equally important partial pressure of oxygen
that it can release oxygen within respiring tissues.
Haemoglobin performs these tasks superbly.

Partial pressure of
1 2 3 4 5 6 7 8 9 10 11 12 13 14
oxygen / kPa
Percentage saturation
8.5 24.0 43.0 57.5 71.5 80.0 85.5 88.0 92.0 94.0 95.5 96.5 97.5 98.0
of haemoglobin

Table 8.2: Effect of oxygen partial pressure on the percentage saturation of haemoglobin with oxygen.

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

The shape of the haemoglobin dissociation curve reflects

Saturation of haemoglobin with oxygen / %

100 the way that oxygen atoms combine with haemoglobin

molecules. Up to an oxygen partial pressure of around
80 2 kPa, on average only one oxygen molecule is combined
with each haemoglobin molecule. Once this oxygen
60 molecule is combined, however, it becomes successively
easier for the second and third oxygen molecules to
combine, so the curve rises very steeply. Over this part
40
of the curve, a small change in the partial pressure of
oxygen causes a very large change in the amount of
20 oxygen which is carried by the haemoglobin.

0
0 2 4 6 8 10 12 14 Questions
Partial pressure of oxygen / kPa
12 Use the dissociation curve in Figure 8.17 to answer
Figure 8.17: The haemoglobin dissociation curve. these questions.
a i The partial pressure of oxygen in the alveoli
Think about the haemoglobin inside a red blood cell of the lungs is about 12 kPa. What is the
in a capillary in the lungs. Here, the partial pressure of percentage saturation of haemoglobin in
oxygen is high, so this haemoglobin is 95–97% saturated the capillaries in the lungs?
with oxygen. This means that almost every haemoglobin ii If 1 g of fully saturated haemoglobin is
molecule is combined with its full complement of eight combined with 1.3 cm3 of oxygen, how
oxygen atoms. much oxygen will 1 g of haemoglobin in the
capillaries in the lungs be combined with?
In an actively respiring muscle, however, where the
partial pressure of oxygen is low, the haemoglobin b i The partial pressure of oxygen in an
will be about 20–25% saturated with oxygen. In other actively respiring muscle is about 2 kPa.
words, the haemoglobin is carrying only a quarter of the What is the percentage saturation of
oxygen it is capable of carrying. haemoglobin in the capillaries of such a
muscle?
This means that haemoglobin coming from the lungs
ii How much oxygen will 1 g of haemoglobin
carries a lot of oxygen but, when it reaches a muscle,
in the capillaries of this muscle be
it releases around three-quarters of it. This released
combined with?
oxygen diffuses out of the red blood cell and into the
muscle where it can be used in respiration. 13 With a partner, build up a list of the ways in which
the structure of haemoglobin is related to its
The S-shaped curve function as an oxygen transport molecule in the
The shape of the haemoglobin dissociation curve can be blood. (You may wish to look back at Chapter 2 to
explained by the behaviour of a haemoglobin molecule remind you about the various levels of structure of
as it combines with or loses oxygen molecules. a protein molecule such as haemoglobin, but try to
do this without looking if you can.)
Oxygen molecules combine with the iron atoms in the
Then use your list to write a brief but full
haem groups of a haemoglobin molecule. You will
explanation of how the structure of haemoglobin
remember that each haemoglobin molecule has four
helps it to carry out its functions.
haem groups. When an oxygen molecule combines with
one haem group, the whole haemoglobin molecule is The Bohr shift
slightly distorted (its 3D shape changes). The shape The behaviour of haemoglobin in picking up oxygen at
change makes it easier for a second oxygen molecule to the lungs, and readily releasing it when in conditions of
combine with a second haem group. This in turn makes low oxygen partial pressure, is exactly what is needed.
it easier for a third oxygen molecule to combine with a But, in fact, haemoglobin is even better at this than is
third haem group. It is then even easier for the fourth shown by the dissociation curve in Figure 8.17. This is
and final oxygen molecule to combine. because the amount of oxygen the haemoglobin carries

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 8 Transport in mammals

is affected not only by the partial pressure of oxygen but solution, haemoglobin helps to maintain the pH of
also by the partial pressure of carbon dioxide. the blood close to neutral. It is acting as a buffer.
Carbon dioxide is continually produced by respiring • The presence of a high partial pressure of carbon
cells. It diffuses from the cells and into blood plasma, dioxide causes haemoglobin to release oxygen. This
from where some of it diffuses into the red blood cells. is called the Bohr shift, after Christian Bohr who
discovered it in 1904. It is exactly what is needed.
In the cytoplasm of red blood cells there is an enzyme,
High concentrations of carbon dioxide are found
carbonic anhydrase, that catalyses the following reaction:
in actively respiring tissues, which need oxygen;
carbonic these high carbon dioxide concentrations cause
anhydrase haemoglobin to release its oxygen even more readily
than it would otherwise do.
CO2 + H2O H2CO3

carbon dioxide water carbonic acid KEY WORDS

carbonic anhydrase: an enzyme found in the
The carbonic acid dissociates: cytoplasm of red blood cells that catalyses the
H2CO3 H+ + HCO3− reaction between carbon dioxide and water to
form carbonic acid
carbonic acid hydrogen ion hydrogencarbonate ion
Bohr shift: the decrease in affinity of
Haemoglobin readily combines with the hydrogen ions, haemoglobin for oxygen that occurs when
forming haemoglobinic acid, HHb. When haemoglobin carbon dioxide is present
does this, it releases the oxygen which it is carrying.
The net result of this reaction is twofold. If a dissociation curve is drawn for haemoglobin at a
• Haemoglobin removes excess hydrogen ions from high partial pressure of carbon dioxide, it looks like the
solution. When carbon dioxide dissolves and lower curve shown on both graphs in Figure 8.18. At
dissociates, a high concentration of hydrogen ions each partial pressure of oxygen, the haemoglobin is less
is formed. This produces a low pH. If the hydrogen saturated than it would be at a low partial pressure of
ions were left in solution, the blood would be carbon dioxide. The curve therefore lies below, and to
very acidic. By removing the hydrogen ions from the right of, the ‘normal’ curve.

The effect of changes in carbon dioxide concentration on The effect of changes in carbon dioxide concentration on
haemoglobin saturation oxygen transport
Saturation of haemoglobin with oxygen / %

Saturation of haemoglobin with oxygen / %

100 100

80 low CO2 80 In lung capillaries, this

concentration
section of this curve is
the relevant one.
60 high CO2 60
concentration
40 40
In active tissues such as
contracting skeletal muscle,
20 20 this section of this curve is
the relevant one.
0 0
0 2 4 6 8 10 12 14 0 2 4 6 8 10 12 14
Partial pressure of oxygen / kPa Partial pressure of oxygen / kPa

Figure 8.18: Dissociation curves for haemoglobin at two different partial pressures of carbon dioxide. The shift of the curve to
the right when the haemoglobin is exposed to higher carbon dioxide concentration is called the Bohr shift.

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

The chloride shift Carbon dioxide transport

The hydrogencarbonate ions that are produced inside Carbon dioxide is produced all over the body by
red blood cells, as a result of the action of carbonic respiring cells. Blood transports this waste product from
anhydrase on carbon dioxide, diffuse out of the cells and the cells to the lungs, where it is lost by diffusion into
into the blood plasma. These ions have a negative charge the alveoli (Chapter 9, Section 9.5, Alveoli). The blood
and, to balance their movement, chloride ions (which also transports carbon dioxide in three different ways.
have a negative charge) move from the blood plasma into
the red blood cells. This is called the chloride shift. As hydrogencarbonate ions in the blood plasma
If the chloride shift did not happen, the inside of The description of the Bohr shift above explains one
the red blood cell would develop an overall positive way in which carbon dioxide is carried in the blood. One
charge, because hydrogen ions (from the dissociation product of the dissociation of dissolved carbon dioxide
of carbonic acid) would accumulate. Hydrogen ions is hydrogencarbonate ions, HCO3–. These are formed
cannot leave the cell, because its cell membrane is not in the cytoplasm of the red blood cell, because this is
permeable to them. The influx of chloride ions therefore where the enzyme carbonic anhydrase is found. Most
helps to prevent the overall charge inside the cell from of the hydrogencarbonate ions then diffuse out of the
becoming too positive. red blood cell into the blood plasma, where they are
carried in solution. About 85% of the carbon dioxide
transported by the blood is carried in this way.
KEY WORDS
chloride shift: the movement of chloride ions As dissolved carbon dioxide molecules in the
into red blood cells from blood plasma, to blood plasma
balance the movement of hydrogencarbonate Some carbon dioxide remains as carbon dioxide
ions into the plasma from the red blood cells molecules and some of these simply dissolve in the blood
plasma. About 5% of the total is carried in this form.
carbaminohaemoglobin: a compound formed
when carbon dioxide binds with haemoglobin As carbaminohaemoglobin
Other carbon dioxide molecules diffuse into the red
blood cells but do not undergo the reaction catalysed
tissue fluid by carbonic anhydrase. Instead, they combine directly
from respiring to alveolar
cells air
with the terminal amine groups (–NH2) of some of
the haemoglobin molecules. The compound formed is
CO2 called carbaminohaemoglobin. About 10% of the carbon
dioxide is carried in this way (Figure 8.19).

plasma 5% CO2
in solution
CO2 + water CO2 +
haemoglobin
carbonic
anhydrase

H+ + HCO−3 H2CO3 carbamino-

haemoglobin
85% hydrogen-
carbonate 10% carbamino-
haemoglobin

red blood cell

Figure 8.19: Carbon dioxide transport in the blood. The blood carries carbon dioxide partly as undissociated carbon dioxide
in solution in the plasma, partly as hydrogencarbonate ions in solution in the plasma, and partly combined with haemoglobin
in the red blood cells.

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 8 Transport in mammals

When blood reaches the lungs, the reactions described

above go into reverse. As there is a relatively low
concentration of carbon dioxide in the alveoli compared
with that in the blood, carbon dioxide diffuses from the
blood into the air in the alveoli. In turn, this stimulates the
carbon dioxide of carbaminohaemoglobin to leave the red
blood cell, and hydrogencarbonate and hydrogen ions to
recombine to form carbon dioxide molecules once more.
This leaves the haemoglobin molecules free to combine
with oxygen, ready to begin another circuit of the body.

Question
14 The following statements were all made by
candidates in examination answers. Explain what is
wrong with each of the following statements.
a Oxyhaemoglobin gradually releases its oxygen
as it passes from the lungs to a muscle.
b The strong walls of arteries enable them to
pump blood around the body.
Figure 8.20: A human heart. The blood vessels in the
c Each red blood cell can combine with eight
photograph lie immediately below the surface of the heart
oxygen atoms.
and have been injected with gelatine containing a dye. The
d Red blood cells have a large surface area so
cardiac muscle was treated to make it transparent to a depth
that many oxygen molecules can be attached.
of 2 mm to allow the blood vessels to be seen.
e Most carbon dioxide is transported in solution
in blood plasma.

aorta
8.6 The heart vena cava left
from head pulmonary
The heart of an adult human has a mass of around
300 g and is about the size of your fist (Figure 8.20). It is artery
right
a bag made of muscle and filled with blood. Figure 8.21 pulmonary left atrium
shows the appearance of a human heart, looking at it artery
from the front of the body. left
right pulmonary
The muscle of which the heart is made is called cardiac
pulmonary vein
muscle. Although you do not need to know the structure vein
of cardiac muscle, you may find it interesting, and it is
left
shown in Figure 8.22. It is made of interconnecting cells ventricle
with cell surface membranes very tightly joined together. right atrium
This close contact between the muscle cells allows waves
of electrical excitation to pass easily between them. This
right ventricle coronary
is a very important feature of cardiac muscle, as you will
arteries
see later.
vena cava from
KEY WORD lower regions of
body
cardiac muscle: the type of muscle that makes
up the walls of the heart Figure 8.21: Diagram of the external structure of a human
heart, seen from the front.

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

KEY WORDS
coronary arteries: arteries that branch from the
aorta and spread over the walls of the heart,
supplying the cardiac muscle with nutrients
and oxygen
septum: the layer of tissue that separates the left
and right sides of the heart
atrium (plural: atria): one of the chambers of
the heart that receives low-pressure blood from
the veins
ventricle: one of the chambers of the heart that
Figure 8.22: Diagram of the structure of cardiac muscle. receives blood from the atria and then pushes it
You do not need to know this structure, but you may like into the arteries
to compare it with striated muscle, shown in Chapter 15 atrioventricular valve: a valve between the atria
(Figure 15.26). and ventricles that closes when the ventricles
contract and stops backflow of blood into the atria
On the surface of the heart, the coronary arteries can bicuspid valve: the atrioventricular valve on the
be seen (Figure 8.20). These branch from the aorta, and left side of the heart
deliver oxygenated blood to the walls of the heart itself.
tricuspid valve: the atrioventricular valve on the
If the heart is cut open vertically (Figures 8.23 and 8.24), it right side of the heart
can be seen to contain four chambers. The two chambers
on the left of the heart are completely separated from
those on the right by a wall of muscle called the septum.
atrium (the
Blood cannot pass through the septum; the only way for space above
blood to get from one side of the heart to the other is the cusp)
for it to leave the heart, circulate around either the lungs tendon
or the rest of the body, and then return to the other side a cusp of one
of the heart. atrioventricular
valve
The upper chamber on each side of the heart is called an papilllary
atrium (plural: atria). The two atria receive blood from muscle
the veins. You can see from Figure 8.24 that blood from
the venae cavae flows into the right atrium, while blood ventricle
from the pulmonary veins flows into the left atrium.
wall of
The lower chambers are ventricles. Blood flows into the ventricle
ventricles from the atria, and is then squeezed out into
the arteries. Blood from the left ventricle flows into the
aorta, while blood from the right ventricle flows into the Figure 8.23: Section through part of the left side of the heart.
pulmonary arteries.
The atria and ventricles have valves between them,
which are known as the atrioventricular valves. The one
on the left is the mitral or bicuspid valve, and the one on
the right is the tricuspid valve.

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 8 Transport in mammals

Figure 8.24: Diagrammatic section through a heart. KEY WORD

cardiac cycle: the
sequence of events that
The cardiac cycle takes place during one
Your heart beats around 70 times a minute. The cardiac cycle is the sequence of heartbeat
events which makes up one heartbeat. Figure 8.25 shows three stages in this cycle.

1 Atrial systole. Both atria contract. 2 Ventricular systole. Both ventricles contract. 3 Ventricular diastole. Atria and ventricles
Blood flows from the atria into the The atrioventricular valves are pushed shut relax. The semilunar valves in the aorta
ventricles. Backflow of blood into by the pressurised blood in the ventricles. and pulmonary artery are pushed shut.
the veins is prevented by closure of The semilunar valves in the aorta and Blood flows from the veins through the
the valves in the veins. pulmonary artery are pushed open. Blood atria and into the ventricles.
flows from the ventricles into the arteries.

pressure exerted by movement of blood

contraction of muscle
Figure 8.25: The cardiac cycle. Only three stages in this continuous process are shown.

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

As the cycle is continuous, a description of it could Ventricular systole lasts for about 0.3 seconds. The
begin anywhere. Let’s begin with the time when the heart muscle then relaxes, and the stage called ventricular
is filled with blood and the muscle in the atrial walls diastole begins. As the muscle relaxes, the pressure in
contracts. This stage is called atrial systole (pronounced the ventricles drops. The high-pressure blood which
sis-tole-ee). The pressure developed by this contraction has just been pushed into the arteries would flow back
is not very great because the muscular walls of the atria into the ventricles, but the semilunar valves snap shut
are only thin, but it is enough to force the blood in the as the blood fills their cusps and so prevent this from
atria down through the atrioventricular valves into the happening.
ventricles. The blood from the atria does not go back
into the pulmonary veins or the venae cavae because
these veins have semilunar valves to prevent backflow. KEY WORDS
About 0.1 seconds after the atria contract, the ventricles atrial systole: the stage of the cardiac cycle when
contract. This is called ventricular systole. The thick, the muscle in the walls of the atria contracts
muscular walls of the ventricles squeeze inwards on ventricular systole: the stage of the cardiac cycle
the blood, increasing its pressure and pushing it out when the muscle in the walls of the ventricles
of the heart. As soon as the pressure in the ventricles contracts
becomes greater than the pressure in the atria, the
pressure difference pushes the atrioventricular valves diastole: the stage of the cardiac cycle when the
shut, preventing blood from going back into the atria. muscle in the walls of the heart relaxes
Instead, the blood rushes upwards into the aorta and the
pulmonary artery, pushing open the semilunar valves in
these vessels as it does so.

Atrioventricular valve
During atrial systole, the pressure of the blood atrial systole –
is higher in the atrium than in the ventricle, and valve open
so forces the atrioventricular valve open.
During ventricular systole, the pressure of the ventricular
blood is higher in the ventricle than in the systole –
atrium. The pressure of the blood pushes up valve shut
against the cusps of the atrioventricular valve,
pushing it shut. Contraction of the papillary
muscles, attached to the valve by tendons,
cusp of valve
prevents the atrioventricular valve from being
forced inside-out.
papillary muscle
tendon
Semilunar valve
During ventricular systole, the pressure of ventricular cusp of valve
the blood forces the semilunar valves open. diastole –
During ventricular diastole, the pressure of valve shut
the blood in the arteries is higher than in the
ventricular
ventricles. The pressure of the blood pushes
systole –
into the cusps of the semilunar valves,
valve open
squeezing them shut.

Figure 8.26: How the heart valves function.

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 8 Transport in mammals

During diastole, as all the heart muscle relaxes, blood that the left ventricle is capable of producing would
from the veins flows into the two atria. The blood is at a be too great. So, as you have seen, arterioles play an
very low pressure, but the thin walls of the atria are easily important role in reducing this pressure before blood
distended, providing very little resistance to the blood flows into the capillaries. However, during vigorous
flow. Some of the blood trickles downwards into the exercise, when muscles are working hard, the arterioles
ventricles, through the atrioventricular valves. The atrial supplying blood to them dilate, increasing blood flow to
muscle then contracts, to push blood forcefully down them. The left ventricle must be able to develop enough
into the ventricles, and the whole cycle begins again. force to ensure that there is still sufficient blood reaching
other organs. For this reason, the thickness of the
Figure 8.26 shows how the atrioventricular and
muscular wall of the left ventricle is much greater than
semilunar valves work. Figure 8.27 shows the pressure
that of the right.
changes in the left side of the heart during one
cardiac cycle. Figure 8.27 shows the pressure changes in the left side of
the heart and the aorta during two consecutive cardiac
The walls of the ventricles are much thicker than the
cycles. You can see that the pressure developed in the left
walls of the atria because the ventricles need to develop
ventricle is much greater than that in the left atrium.
much more force when they contract. Their contraction
has to push the blood out of the heart and around the
body. For the right ventricle, the force produced must
be relatively small, because the blood goes only to the Questions
lungs, which are very close to the heart. If the pressure 15 From Figure 8.27, identify the time at which each
developed was too high, lung capillaries could be stage shown in Figure 8.25 is occurring.
damaged and tissue fluid would accumulate in the lungs, 16 Heart valves can become weakened and fail to
hampering gas exchange. close effectively. Suggest how this would affect the
The left ventricle, however, has to develop sufficient function of the heart and the health of a person.
force to supply blood to all the rest of the body organs.
For most organs, most of the time, the high pressures

16

14
aortic pressure
12

10
Pressure / kPa

8
left ventricular
pressure
6

4
left atrial
2 pressure

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1 1.2 1.3
Time / s

Figure 8.27: Pressure changes in the left side of the heart during the cardiac cycle.

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Control of heartbeat walls. The cardiac muscle in the atrial walls responds to
this excitation wave by contracting, at the same rhythm
Cardiac muscle differs from the muscle in all other as the SAN. Therefore, all the muscle in both atria
areas of the body in that it is myogenic. This means that contracts almost simultaneously.
it naturally contracts and relaxes; it does not need to
As you have seen, the muscles of the ventricles do not
receive impulses from a nerve to make it contract. If
contract until after the muscles of the atria. (You can
cardiac muscle cells are cultured in a warm, oxygenated
imagine what would happen if they all contracted at
solution containing nutrients, they contract and relax
once.) This delay is caused by a feature of the heart that
rhythmically, all by themselves.
briefly delays the excitation wave in its passage from the
However, the individual heart muscle cells cannot be atria to the ventricles. There is a band of fibres between
allowed to contract at their own natural rhythms. If the atria and ventricles which does not conduct the
they did, parts of the heart would contract out of excitation wave. This means that, as the wave spreads
sequence with other parts. The cardiac cycle would out from the SAN over the atrial walls, it cannot pass
become disordered, and the heart would stop working as directly into the ventricle walls. The only route through
a pump. The heart has its own built-in controlling and is via a patch of conducting fibres, situated in the
coordinating system which prevents this happening.
The cardiac cycle is initiated in a specialised patch KEY WORDS
of muscle in the wall of the right atrium called the myogenic: a word used to describe muscle tissue
sinoatrial node. It is often called the SAN for short, or that contracts and relaxes even when there is no
pacemaker. The muscle cells of the SAN set the rhythm stimulation from a nerve
for all the other cardiac muscle cells. Their natural
rhythm of contraction is slightly faster than that of the sinoatrial node (SAN): a patch of cardiac muscle
rest of the heart muscle. Each time the muscles of the in the right atrium of the heart which contracts
SAN contract, they set up a wave of electrical activity and relaxes in a rhythm that sets the pattern for
which spreads out rapidly over the whole of the atrial the rest of the heart muscle

Figure 8.28: How electrical excitation waves move through the heart.

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 8 Transport in mammals

septum, known as the atrioventricular node, or AVN

(Figure 8.28). The AVN picks up the excitation wave as KEY WORDS
it spreads across the atria and, after a delay of about atrioventricular node (AVN): a patch of tissue
0.1 seconds, passes it on to a bunch of conducting fibres in the septum of the heart which transmits the
called the Purkyne tissue, which runs down the septum wave of excitation from the walls of the atria and
between the ventricles. This transmits the excitation transmits it to the Purkyne tissue
wave very rapidly down to the base of the septum,
from where it spreads outwards and upwards through Purkyne tissue: a bundle of fibres that conduct
the ventricle walls. As it does so, it causes the cardiac the wave of excitation down through the septum
muscle in these walls to contract, from the bottom up, so of the heart to the base (apex) of the ventricles
squeezing blood upwards and into the arteries.

REFLECTION
In your group, discuss the transport systems of flowering plants and mammals.
• What functions do they have in common, and how do the structures of each of them help to fulfil these
functions?
• How do the differences between them reflect the differences in the requirements of plants and animals?
What process did you go through to produce your responses?

Final reflection
Discuss with a friend which, if any, part of Chapter 8 you need to:
• read through again to make sure you really understand
• seek more guidance on, even after going over it again.

SUMMARY

Blood is carried away from the heart in arteries, passes through tissues in capillaries, and is returned to the heart
in veins. Blood pressure drops gradually as it passes along this system.
Arteries have thick, elastic walls, to allow them to withstand high blood pressures and to smooth out the pulsed
blood flow. Arterioles are small arteries that help to reduce blood pressure and control the amount of blood
flow to different tissues. Capillaries are only just wide enough to allow the passage of red blood cells, and
they have very thin walls to allow efficient and rapid transfer of materials between blood and cells. Veins have
thinner walls than arteries and possess valves to help blood at low pressure flow back to the heart.
Blood plasma leaks from capillaries to form tissue fluid.
Red blood cells are relatively small cells. They have a biconcave shape and no nucleus. Their cytoplasm is full of
haemoglobin.
White blood cells include phagocytes and lymphocytes. They all have nuclei, and are either spherical or
irregular in shape.
Red blood cells carry oxygen in combination with haemoglobin.

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

CONTINUED

Haemoglobin picks up oxygen at high partial pressures (concentrations) of oxygen in the lungs and releases it at
low partial pressures of oxygen in respiring tissues. A graph showing the percentage saturation of haemoglobin
at different partial pressures of oxygen is known as a dissociation curve.
At high carbon dioxide concentrations, the dissociation curve shifts downwards and to the right, showing that
haemoglobin releases oxygen more easily when carbon dioxide concentration is high. This is known as the Bohr shift.
The mammalian heart has four chambers: right and left atria and right and left ventricles. The right side of the
heart is divided from the left by a wall of tissue called the septum. The atrial muscular walls are thin and do not
exert much pressure when they contract. The ventricular walls are much more muscular and exert a sufficient
pressure to drive blood to the lungs from the right ventricle and around the rest of the body from the left
ventricle. The left ventricular wall is therefore much thicker and more muscular than the right ventricular wall.
The cardiac cycle is a continuous process but can be considered in three stages. 1 Atrial systole (contraction of
the atria) allows blood to flow into the ventricles from the atria. Closure of valves in the veins prevents backflow
of blood into the veins. 2 Ventricular systole (contraction of the ventricles) pushes blood into the arteries by
forcing open the semilunar valves. Blood is prevented from flowing back into the atria by pressure closing the
atrioventricular valves. 3 In diastole (relaxation of heart muscle), the semilunar valves are pushed shut, preventing
backflow of blood from the arteries into the ventricles. Blood flows into the atria and ventricles from the veins.
Beating of the heart is initiated by the sinoatrial node (SAN) or pacemaker, which has its own myogenic rhythm.
A wave of excitation spreads across the atria so all the heart muscle cells in the atria contract together. The wave
of excitation cannot spread to the ventricles directly because of a band of non-conducting tissue. However, the
atrioventricular node (AVN) in the septum passes the wave to the Purkyne tissue, which then causes the ventricles
to contract from the bottom up shortly after the atria. This is important because it pushes the blood upwards out
of the ventricles into the arteries.

EXAM-STYLE QUESTIONS

1 Where is the mammalian heartbeat initiated?

A atrioventricular node (AVN)
B left atrium
C Purkyne tissue
D sinoatrial node (SAN) [1]
2 What causes the bicuspid valve to close during ventricular systole?
A a greater blood pressure in the left atrium than in the left ventricle
B a greater blood pressure in the left ventricle than in the left atrium
C contraction of muscles in the septum
D contraction of muscles in the valve [1]

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 8 Transport in mammals

CONTINUED
3 The diagram shows the changes in blood pressure as blood flows through the
blood vessels in the human systemic circulatory system.
Which correctly identifies the vessels labelled P to S?

20

15
Pressure / kPa

10

P Q R S
5

0
Blood vessels (direction of blood flow →)

P Q R S
A artery capillary arteriole venule
B arteriole artery venule capillary
C artery arteriole capillary venule
D venule capillary arteriole artery
[1]
4 The micrograph shows an artery and a vein.

X Y

Which row correctly identifies and describes the artery and the vein?
X Y Description
A artery vein The artery has thick walls and the vein has thin walls.
B artery vein The artery has a thin middle layer while the
vein has a thick middle layer.
C vein artery The artery has a thick middle layer while the
vein has a thin middle layer.
D vein artery The artery has thin walls and the vein has thick walls. [1]

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

CONTINUED
5 Carbon dioxide is transported in the blood in various forms.
a Describe how carbon dioxide molecules reach red blood cells from
respiring cells. [2]
b The diagram below shows part of a capillary network and some cells of
the surrounding tissue.

arteriole end venule end

X Y
State three ways in which the blood at Y differs from the blood
at X other than in the concentration of carbon dioxide. [3]
c An enzyme in red blood cells catalyses the reaction between carbon dioxide
and water as blood flows through respiring tissues.
enzyme
CO2 + H2O H2CO3 H+ + HCO3−

i Name the enzyme that catalyses this reaction. [1]

ii Explain the significance of this reaction in the transport of
carbon dioxide. [3]
d The graph below shows the effect of increasing the carbon dioxide
concentration on the oxygen dissociation curve for haemoglobin.
100
Saturation of haemoglobin with

80 partial pressure
CO2 1.0 kPa
oxygen / %

60
partial pressure
CO2 1.5 kPa
40

20

0
0 2 4 6 8 10 12 14
Partial pressure of oxygen / kPa

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 8 Transport in mammals

CONTINUED
i State the percentage saturation of haemoglobin with oxygen at a
partial pressure of 5 kPa of oxygen when the partial pressure of
carbon dioxide is:
1.0 kPa
1.5 kPa [1]
ii The percentage saturation of haemoglobin with oxygen decreases as the
partial pressure of carbon dioxide increases. Explain how this happens.
[2]
iii Name the effect of increasing carbon dioxide concentration on the
oxygen dissociation curve. [1]
iv Explain the importance of the effect of carbon dioxide on haemoglobin
as shown in the graph. [3]
[Total: 16]
Cambridge International AS & A Level Biology (9700) Paper 21, Question 2,
June 2011

6 Mammals have a closed, double circulation.

a State what is meant by the term double circulation. [1]
b The figure below shows part of the circulation in a mammalian tissue.
The central part is enlarged to show a capillary and a cell supplied by
the capillary.

pre-capillary sphincter
muscle

vein
artery
plasma
arteriole
tissue fluid

Explain why the wall of the artery is thicker than the wall of the vein. [2]
c Suggest one role for the pre-capillary sphincter muscle shown in
the figure. [1]
d With reference to the figure, describe the role of capillaries in forming
tissue fluid. [3]
e Describe three ways in which plasma differs from tissue fluid. [3]
[Total: 10]
Cambridge International AS & A Level Biology (9700) Paper 2, Question
4a–ei, November 2008

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

CONTINUED
7 Figure 8.27 shows the pressure changes in the left atrium, left ventricle and
aorta throughout two cardiac cycles. Make a copy of this diagram.
a i How long does one heartbeat (one cardiac cycle) last? [1]
ii What is the heart rate represented on this graph, in beats per minute? [1]
b The contraction of muscles in the ventricle wall causes the pressure inside
the ventricle to rise. When the muscles relax, the pressure drops again. On
your copy of the diagram, mark the following periods:
i the time when the ventricle is contracting (ventricular systole) [1]
ii the time when the ventricle is relaxing (ventricular diastole). [1]
c The contraction of muscles in the wall of the atrium raises the pressure
inside it. This pressure is also raised when blood flows into the atrium from
the veins, while the atrial walls are relaxed. On your copy of the diagram,
mark the following periods:
i the time when the atrium is contracting (atrial systole) [1]
ii the time when the atrium is relaxing (atrial diastole). [1]
d The atrioventricular valves open when the pressure of the blood in the atria
is greater than that in the ventricles. They snap shut when the pressure of the
blood in the ventricles is greater than that in the atria.
On your diagram, mark the points at which these valves will open
and close. [1]
e The opening and closing of the semilunar valves in the aorta depends in a
similar way on the relative pressures in the aorta and ventricles. On your
diagram, mark the points at which these valves will open and close. [1]
f The right ventricle has much less muscle in its walls than the left ventricle,
and only develops about one-quarter of the pressure developed on the left
side of the heart. On your diagram, draw a line to represent the probable
pressure inside the right ventricle over the 1.3 seconds shown. [1]
[Total: 9]
8 The diagram below shows a cross section of the heart at the level of the valves.
Q

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 8 Transport in mammals

CONTINUED
a i Copy and complete the following flow chart to show the pathway of
blood through the heart.

vena cava right atrium valve P valve S

left atrium lungs pulmonary artery

valve Q left ventricle valve R aorta

ii Explain how the valves P and Q ensure one-way flow of blood through
the heart. [2]
b The cardiac cycle describes the events that occur during one heartbeat. The
following figure shows the changes in blood pressure that occur within the
left atrium, left ventricle and aorta during one heartbeat. [2]
1 2
16

12
5
Blood pressure / kPa

8 4

4
3 6
7
left atrium
left ventricle
0 aorta
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Time / s

Copy and complete the table below. Match up each event during the cardiac
cycle with an appropriate number from 1 to 7 on the diagram. You should put
only one number in each box. You may use each number once, more than once or
not at all.
The first answer has been completed for you.
Event during the cardiac cycle Number
atrioventricular (bicuspid) valve opens 6
ventricular systole
semilunar (aortic) valve closes
left ventricle and left atrium both relaxing
semilunar (aortic) valve opens

[4]

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 CAMBRIDGE INTERNATIONAL AS & A LEVEL BIOLOGY: COURSEBOOK

CONTINUED

c Explain the roles of the SAN, AVN and the Purkyne tissue during one
heartbeat. [5]
[Total: 13]
Cambridge International AS & A Level Biology (9700) Paper 21,
Question 3, May/June 2010

SELF-EVALUATION CHECKLIST
After studying this chapter, complete a table like this:

See Needs Almost Ready to

I can
section... more work there move on
describe the structure of the mammalian 8.2
circulatory system as a closed, double
circulation, and describe its main components
explain how the structures of arteries, arterioles, 8.3
veins, venules and capillaries are related to their
functions
describe the formation and functions of tissue 8.4
fluid
describe the structure of blood, including blood 8.5
plasma, red cells, monocytes, neutrophils and
lymphocytes
make diagrams of blood vessels and blood 8.4, 8.5
cells from slides, photomicrographs or electron
micrographs
explain, in detail, how oxygen and carbon 8.5
dioxide are transported in the blood, and
interpret oxygen dissociation curves
describe and explain the Bohr shift 8.5

explain how the solvent properties of water, and 8.5

its high heat capacity, enable blood plasma and
tissue fluid to carry out their functions
explain the structure and function of the heart 8.6

describe the cardiac cycle and its control 8.6

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