Notes:

Double circulatory system: one in which the blood flows through the heart twice for each
circuit of the body.
Single circulatory system: one in which the blood flows through the heart once for each
circuit of the body.
Transport: the movement of substances such as oxygen, nutrients, hormones, waste and
heat around the body.

The need for a transport system

All living animal cells need a supply of oxygen and nutrients to grow and survive. They also
need to remove waste products so that these do not build up and become toxic. Very small
animals do not need a separate transport system, because all their cells are surrounded by
(or very close to) the environment in which they live. Diffusion will supply enough oxygen
and nutrients to keep the cell alive. However, a larger animal with a complex anatomy will
have more than two layers of cells. The diffusion distance becomes too long, and diffusion
alone will be too slow to supply all the requirements. There are three main factors that
influence the need for a transport system:
• size
• surface area to volume ratio
• level of metabolic activity.

Size
The cells inside a large organism are further from its surface — the diffusion pathway is
increased. The diffusion rate is reduced, and diffusion is too slow to supply all the
requirements. Also, the outer layers of cells use up the supplies, so that less will reach the
cells deep inside the body.

Surface area to volume ratio

Small animals have a large surface area to volume ratio. This means that for each gram of
tissue in their body they have a sufficient area of the body surface through which exchange
can occur. However, larger animals have a smaller surface area to volume ratio. This means
that each gram of tissue has a smaller area of the body surface for exchange.

Level of metabolic activity Animals need energy from food, so that they can move around.
Releasing energy from food by aerobic respiration requires oxygen. If an animal is very
active, its cells need good supplies of nutrients and oxygen to supply the energy for
movement. Animals that keep themselves warm, such as mammals, need even more
energy.

Features of a good transport system

An effective transport system will include:
• a fluid or medium to carry nutrients, oxygen and wastes around the body — this is the
blood
• a pump to create pressure that will push the fluid around the body — this is the heart
• exchange surfaces that enable substances to enter the blood and leave it again where they
are needed — these are the capillaries.

An efficient transport system will also include:

• tubes or vessels to carry the blood by mass flow
• two circuits — one to pick up oxygen and another to deliver oxygen to the tissues.

Single and double circulatory systems

Fish have a single circulatory system. The blood flows through the heart once for each
circuit of the body. The blood takes the following route: heart->gills-> body->heart

Mammals have a double circulatory system. The system has two separate circuits. One
circuit carries blood to the lungs to pick up oxygen. This is pulmonary circulation. The other
circuit carries the oxygen and nutrients around the body to the tissues. This is systemic
circulation. Blood flows through the heart twice for each circuit of the body. The blood takes
the following route: heart-> body-> heart -> lungs-> heart

Advantages of a double circulation

An efficient circulatory system will deliver oxygen and nutrients quickly to the parts of the
body where they are needed. The blood can be made to flow more quickly by increasing the
blood pressure created by the heart.
In the single circulatory system of fish:
• the blood pressure drops as blood passes through the tiny capillaries of the gills
• blood has a low pressure as it flows towards the body, and will not flow very quickly
• the rate at which oxygen and nutrients are delivered to respiring tissues, and carbon
dioxide and urea are removed, is limited. Fish are not as metabolically active as mammals,
as they do not maintain their body temperature. Therefore, they need less energy. Their
single circulatory system delivers sufficient oxygen and nutrients for their needs.

In the double circulatory system of mammals:

• the blood pressure must not be too high in the pulmonary circulation, otherwise it may
damage the delicate capillaries in the lungs.
• the heart can increase the pressure of the blood after it has passed through the lungs, so
the blood is under higher pressure as it flows to the body and flows more quickly
• the systemic circulation can carry blood at a higher pressure than the pulmonary
circulation.
Mammals are active animals and maintain their body temperature. Supplying the energy for
activity and the heat needed to keep the body warm requires energy from food. The energy
is released from food in the process of respiration. To release a lot of energy, the cells need
a good supply of both nutrients and oxygen, as well as the removal of waste products.

arteries: vessels that carry blood away from the heart

arterioles: small blood vessels that distribute blood from an artery to the capillaries
capillaries: very small vessels with very thin walls
closed circulatory system: one in which blood is held in vessels
open circulatory system: one in which blood is not held in vessels
veins: vessels that carry blood back to the heart
venules: small blood vessels that collect blood from capillaries and lead into the veins

Open circulatory systems

Many animals, including insects, have an open circulatory system. This means that the
blood is not always held within blood vessels. Instead, the blood fluid circulates through the
body cavity, so that the tissues and cells are bathed directly in blood. In some animals,
movements of the body help to circulate the blood — and without movement the blood stops
moving, so that transport of oxygen and nutrients stops. In other animals, such as insects,
there is a muscular pumping organ much like a heart. This is a long, muscular tube that lies
just under the dorsal (upper) surface of the body. Blood from the body enters the heart
through pores called ostia. The heart then pumps the blood towards the head by peristalsis.
At the forward end of the heart (nearest the head), the blood simply pours out into the body
cavity. This circulation can continue when the insect is at rest, but body movements may still
affect circulation.
Some larger and more active insects, such as locusts, have open-ended tubes attached to
the heart. These direct the blood towards active parts of the body, such as the leg and wing
muscles. Open circulatory systems have certain disadvantages:
• Blood pressure is low and blood flow is slow
• Circulation of blood may be affected by body movements or lack of body movements.

Closed circulatory systems

In larger animals the blood stays entirely inside vessels — this is a closed circulatory
system. A separate fluid, called tissue fluid, bathes the tissues and cells. This has certain
advantages over the open system:
• Higher pressure, so that blood flows more quickly
• More rapid delivery of oxygen and nutrients
• More rapid removal of carbon dioxide and other wastes
• Transport is independent of body movements.

Blood vessels
Blood flows through a series of vessels. Each is adapted to its particular role in relation to its
distance from the heart. All types of blood vessel have an inner layer or lining, made of a
single layer of cells, called the endothelium. This is a thin layer that is particularly smooth in
order to reduce friction with the flowing blood.

Arteries
Arteries carry blood away from the heart. The blood is at high pressure, so the artery wall
must be thick in order to withstand that pressure. The lumen is relatively small in order to
maintain high pressure, and the inner wall is folded to allow the lumen to expand as blood
flow increases. The wall consists of three layers:
• Inner layer (tunica intima) consists of a thin layer of elastic tissue which allows the wall to
stretch and then recoil to help maintain blood pressure.
• Middle layer (tunica media) consists of a thick layer of smooth muscle.
• Outer layer (tunica adventitia) is a relatively thick layer of collagen and elastic tissue. This
provides strength to withstand the high pressure, and recoil to maintain the pressure.
Arteries near the heart have more elastic tissue in the wall, in order to allow stretch and
recoil, which helps to even out the fluctuations in blood pressure created by the heart.
Further from the heart, the walls contain more muscle tissue.

Arterioles
Arterioles are small blood vessels that distribute the blood from an artery to the capillaries.
Arteriole walls contain a layer of smooth muscle. Contraction of this muscle will constrict the
diameter of the arteriole. This increases resistance to flow and reduces the rate of flow of
blood. Constriction of the arteriole walls can be used to divert the flow of blood to regions of
the body that are demanding more oxygen.
Capillaries
Capillaries have very thin walls. They allow exchange of materials between the blood and
tissue fluid.
• The lumen is very narrow — its diameter is about the same as that of a red blood cell. The
red blood cells may be squeezed against the walls of the capillary as they pass along the
capillary; this helps transfer of oxygen, as it reduces the diffusion path to the tissues. It also
increases resistance and reduces rate of flow
• The walls consist of a single layer of flattened endothelial cells. This reduces the diffusion
distance for the materials being exchanged.
• The walls are leaky. They allow blood plasma and dissolved substances to leave the
blood.

Venules
From the capillaries blood flows into small vessels called venules. These collect the blood
from the capillary bed and lead into the veins. The venule wall consists of thin layers of
muscle and elastic tissue outside the endothelium, and a thin outer layer of collagen.

Veins
Veins carry blood back to the heart. The blood is at low pressure and the walls do not need
to be thick.
• The lumen is relatively large, in order to ease the flow of blood.
• The walls have thinner layers of collagen, smooth muscle and elastic tissue than in artery
walls. They do not need to stretch and recoil, and are not actively constricted in order to
reduce blood flow
• The main feature of veins is that they contain valves to help the blood flow back to the
heart and to prevent it flowing in the opposite direction. As the walls are thin, the vein can be
flattened by the action of the surrounding skeletal muscle. Contraction of the surrounding
skeletal muscle applies pressure to the blood, forcing the blood to move along in a direction
determined by the valves.

Blood: the fluid used to transport materials around the body

hydrostatic pressure: the pressure that a fluid exerts when pushing against the sides of a
vessel or container
lymph: the fluid held in the lymphatic system, which is a system of tubes that returns excess
tissue fluid to the blood system.
oncotic pressure: the pressure created by the osmotic effects of the solutes.
plasma: the fluid portion of the blood. tissue fluid: the fluid surrounding the cells and tissues.

Blood plasma and tissue fluid

Blood is the fluid held in our blood vessels. It consists of a liquid called plasma, containing
many blood cells. The plasma contains many dissolved substances, including oxygen,
carbon dioxide, minerals, glucose, amino acids, hormones and plasma proteins. The cells
include the red blood cells (erythrocytes), various white blood cells (leucocytes) and
fragments called platelets. Tissue fluid is similar to blood plasma. But it does not contain
most of the cells found in blood, and neither does it contain plasma proteins. Tissue fluid is
formed by plasma leaking from the capillaries. It surrounds the cells in the tissue, and
supplies them with the oxygen and nutrients they require. As blood plasma leaks from the
capillary, it carries all the dissolved substances into the tissue fluid. This movement is mass
flow rather than diffusion. Waste products from cell metabolism will be carried back into the
capillary as some of the tissue fluid returns to the capillary.

Formation of tissue fluid:

• When an artery reaches the tissues, it branches into smaller arterioles, and then into a
network of capillaries. These eventually link up with venules to carry blood back to the veins.
Therefore blood flowing into an organ or tissue is contained in the capillaries.
• At the arterial end of a capillary, the blood is at relatively high (hydrostatic) pressure. This
pressure tends to push the blood fluid out of the capillaries through the capillary wall. The
fluid can leave through the tiny gaps between the cells in the capillary wall.
• The fluid that leaves the blood consists of plasma with dissolved nutrients and oxygen. All
the red blood cells, platelets and most of the white blood cells remain in the blood, as do the
plasma proteins. These are too large to be pushed out through the gaps in the capillary wall.
This tissue fluid surrounds the body cells, so exchange of gases and nutrients can occur
across the plasma membranes. The exchange occurs by diffusion, facilitated diffusion and
active uptake. Oxygen and nutrients enter the cells; carbon dioxide and other wastes leave
the cells.

Returning to the blood

The blood pressure at the venous end of the capillary is much lower. This allows some of
the tissue fluid to return to the capillary carrying carbon dioxide and other waste substances
into the blood.
Not all of all the tissue fluid re-enters the blood. Some tissue fluid is directed into another
tubular system called the lymph system or lymphatic system. This drains excess tissue fluid
out of the tissues and returns it to the blood system in the subclavian vein in the chest.

The fluid in the lymphatic system is called lymph and is similar in composition to the tissue
fluid. It will contain more lymphocytes, as these are produced in the lymph nodes. Lymph
nodes are swellings found at intervals along the lymphatic system, which have an important
part to play in the immune response.
If a tissue is infected, then the capillaries become more leaky and more fluid is directed into
the lymph system — this helps direct bacteria towards the Lymph nodes.
Blood plasma has high hydrostatic pressure, more negative oncotic pressure than tissue
fluid, and contains red blood cells, neutrophils, and lymphocytes. It also contains plasma
proteins and fats transported in lipoproteins.
Tissues fluid has a low hydrostatic pressure and a less negative oncotic pressure than blood
plasma. It contains some neutrophils, especially in infected areas. It contains few proteins
and few fats.
Lymph has a low hydrostatic pressure. It has a less negative oncotic pressure than blood
plasma and contains lymphocytes, few proteins, and more fats than in tissue fluid, especially
near the digestive system.

Movement of fluids
The hydrostatic pressure of the blood is not the only influence on the movement of fluid into
and out of the capillary. The tissue fluid has its own hydrostatic pressure, and the oncotic
pressure of the solutes (dissolved substances) also has an influence.

• The hydrostatic pressure of the blood tends to push fluid out into the tissues.
• The hydrostatic pressure of the tissue fluid tends to push fluid into the capillaries.
• The oncotic pressure of the blood tends to pull water back into the blood (it has a negative
figure).
• The oncotic pressure of the tissue fluid pulls water into the tissue fluid.
The net result of these forces creates a pressure gradient to push fluid out of the capillary at
the arterial end and into the capillary at the venule end.

atrioventricular valves: valves between the atria and the ventricles, which ensure that blood
flows in the correct direction
cardiac muscle: specialised muscle found in the walls of the heart chambers
semilunar valves: valves that prevent blood re-entering the heart from the arteries.

The mammalian heart is a muscular pump. It is divided into two sides. The right side pumps
the deoxygenated blood to the lungs to be oxygenated. The left side pumps oxygenated
blood to the rest of the body. On both sides, the heart squeezes the blood, putting it under
pressure. This pressure forces the blood along the arteries and through the circulatory
system.

External features of the heart

In humans, the heart lies just off-centre towards the left of the chest cavity. The main part of
the heart consists of firm, dark-red muscle called cardiac muscle. There are two main
pumping chambers — the ventricles. Above the ventricles are two thin-walled chambers —
the atria. These are much smaller than the ventricles and are easy to overlook. Lying over
the surface of the heart are coronary arteries that supply oxygenated blood to the heart
muscle. As the heart is a hard-working organ, these arteries are very important. If they
become constricted, it can have severe consequences for the health of the heart. Restricted
blood flow to the heart muscle reduces the delivery of oxygen and nutrients such as fatty
acids and glucose. This may cause angina or a heart attack (myocardial infarction).

At the top of the heart are a number of tubular blood vessels. These are the veins that carry
blood into the atria and the arteries that carry blood away from the heart.

Internal features of the mammalian heart

The heart is divided into four chambers. The two upper chambers are atria. These receive
blood from the major veins. Deoxygenated blood from the body flows through the vena cava
into the right atrium. Oxygenated blood from the lungs flows through the pulmonary vein into
the left atrium. From the atria, blood flows down through the atrioventricular valves into the
ventricles. Attached to the valves are tendinous cords, which prevent the valves from turning
inside out when the ventricle walls contract. A wall of muscle called the septum separates
the ventricles from each other. This ensures that the oxygenated blood in the left side of the
heart and the deoxygenated blood in the right side are kept separate. Deoxygenated blood
leaving the right ventricle flows into the pulmonary artery leading to the lungs, where it is
oxygenated. Oxygenated blood leaving the left ventricle flows into the aorta. This carries
blood to a number of arteries that supply all parts of the body. At the base of the major
arteries, where they exit the heart, are the semilunar valves. These prevent blood returning
to the heart as the ventricles relax.

Blood pressure
The cardiac muscle in the wall of each chamber contracts to create pressure in the blood.
The higher the pressure created in the heart, the further it will push the blood.
• Atria — the muscle of the atrial walls is very thin. This is because these chambers do not
need to create much pressure. Their function is to receive blood from the veins and push it
into the ventricles.
• Right ventricle — the walls of the right ventricle are thicker than the walls of the atria. This
enables the right ventricle to pump blood out of the heart. The right ventricle pumps
deoxygenated blood to the lungs. The lungs are in the chest cavity beside the heart, so that
the blood does not need to travel very far. Also, the alveoli in the lungs are very delicate and
could be damaged by very high blood pressure.
• Left ventricle — the walls of the left ventricle can be two or three times thicker than those of
the right ventricle. The blood from the left ventricle is pumped out through the aorta and
needs sufficient pressure to overcome the resistance of the systemic circulation.

Cardiac muscle consists of fibres that branch, producing cross-bridges. These help to
spread the stimulus around the heart, and also ensure that the muscle can produce a
squeezing action rather than a simple reduction in length. There are numerous mitochondria
between the muscle fibres (myofibrils) to supply energy for contraction. The muscle cells are
separated by intercalated discs, which facilitate synchronised contraction. Each cell has a
nucleus and is divided into contractile units called sarcomeres.

cardiac cycle: the sequence of events in one full beat of the heart.
In atrial systole both the right and left atria contract together. The muscle in the walls is thin
so only a small increase in pressure is created by this contraction which helps to push blood
into the ventricles stretching their walls and ensuring that they are full of blood. In ventricular
systole both the right and left ventricles pump together. Contraction starts at the apex (base)
of the heart so that blood is pushed upwards towards the arteries.
In diastole the muscular walls of all four chambers relax. Elastic recoil causes the chambers
to increase in volume allowing blood to flow in from the veins.

The action of the valves

The valves ensure that blood flows in the correct direction. They are opened and closed by
changes in the blood pressure in the various chambers of the heart.
Atrioventricular valves
After systole, the ventricular walls relax and recoil.
• The pressure in the ventricles rapidly drops below the pressure in the atria.
• Blood in the atria pushes the atrioventricular valves open.
• Blood entering the heart flows straight through the atria and into the ventricles.
• The pressure in the atria and the ventricles rises slowly as they fill with blood.
• The valves remain open while the atria contract, but close when the atria begin to relax. •
This closure is caused by a swirling action in the blood around the valves when the ventricle
is full.
• As the ventricles begin to contract (systole), the pressure of the blood in the ventricles
rises.
• When the pressure rises above that in the atria, the blood starts to move upwards. • This
movement fills the valve pockets and keeps them closed. • The tendinous cords attached to
the valves prevent them from turning inside out. • This prevents the blood flowing back into
the atria.

Semilunar valves
Before ventricular contraction, the pressure in the major arteries is higher than the pressure
in the ventricles. This means that the semilunar valves are closed. Ventricular systole raises
the blood pressure in the ventricles very quickly. Once the pressure in the ventricles rises
above the pressure in the major arteries, the semilunar valves are pushed open. The blood
is under very high pressure, so it is forced out of the ventricles in a powerful spurt. Once the
ventricle walls have finished contracting, the heart muscle starts to relax (diastole). Elastic
tissue in the walls of the ventricles recoils. This stretches the muscle out again and returns
the ventricle to its original size. This causes the pressure in the ventricles to drop quickly As
it drops below the pressure in the major arteries, the blood starts to flow back towards the
ventricles. The semilunar valves are pushed closed by the blood collecting in the pockets of
the valves. This prevents blood returning to the ventricles. The pressure wave created when
the left semilunar valve closes is the 'pulse' that we can easily feel at the wrist or neck.

pressure changes in the heart chambers

The pressure changes in the heart chambers cause the opening and closing of the valves.

Pressure in the blood vessels

Blood enters the aorta and pulmonary artery in a rapid spurt, but the tissues require blood to
be delivered in an even flow. The structure of the artery walls plays a large part in creating a
more even flow:
• The artery walls close to the heart have a lot of elastic tissue.
• When blood leaves the heart, these walls stretch.
• As blood moves on and out of the aorta, the pressure in the aorta starts to drop.
• The elastic recoil of the walls helps to maintain the blood pressure in the aorta.
• The further the blood flows along the arteries, the more the pressure drops and the
fluctuations become less obvious.
• It is important to maintain the pressure gradient between the aorta and the arterioles, as
this is what keeps the blood flowing towards the tissues.

Coordination of the cardiac cycle

bradycardia: a slow heart rhythm
ectopic heartbeat: an extra beat or an early beat of the ventricles
electrocardiogram: a trace that records the electrical activity of the heart
fibrillation: uncoordinated contraction of the atria and ventricles
myogenic muscle: muscle that can initiate its own contraction
Purkyne tissue: consists of specially adapted muscle fibres that conduct the wave of
excitation from the AVN down the septum to the ventricles
sinoatrial node (SAN): the heart's pacemaker. It is a small patch of tissue that sends out
waves of electrical excitation at regular intervals in order to initiate contractions
tachycardia: a rapid heart rhythm.

The need for coordination

Heart muscle or cardiac muscle is unusual in that it can initiate its own contraction. Because
of this property, the heart muscle is described as myogenic. The muscle will contract and
relax rhythmically even if it is not connected to the body. The muscles from the atria and the
muscles from the ventricles each have their own natural frequency of contraction. The atrial
muscle tends to contract at a higher frequency than the ventricular muscle. This property of
the muscle could cause inefficient pumping if the contractions of the chambers are not
synchronised — a condition known as fibrillation. So the heart needs a mechanism that can
coordinate the contractions of all four chambers.

Initiation and control of the heartbeat

At the top of the right atrium, near the point where the vena cava empties blood into the
atrium, is the sinoatrial node (SAN). This is a small patch of tissue that generates electrical
activity. The SAN initiates a wave of excitation at regular intervals. In a human, this occurs
55-80 times a minute. The SAN is also known as the pacemaker.

Contraction of the atria

• The wave of excitation quickly spreads over the walls of both atria.
• It travels along the membranes of the muscle tissue.
• As the wave of excitation passes, it causes the cardiac muscle cells to contract.
• This is an atrial systole.
The tissue at the base of the atria is unable to conduct the wave of excitation, and so it
cannot spread directly down to the ventricle walls. At the top of the interventricular septum
(the septum separating the two ventricles) is another node — the atrioventricular node
(AVN). This is the only route that can conduct the wave of excitation through to the
ventricles. The wave of excitation is delayed in the node. This allows time for the atria to
finish contracting and for the blood to flow down into the ventricles before they begin to
contract.

Contraction of the ventricles

• After this short delay the wave of excitation is carried away from the AVN and down
specialised conducting tissue called the Purkyne tissue.
• This runs down the interventricular septum
• At the base of the septum, the wave of excitation spreads out over the walls of the
ventricles.
• As the excitation spreads upwards from the base (apex) of the ventricles, it causes the
muscles to [Link] means that the ventricles contract from the base upwards.
• This pushes the blood up towards the major arteries at the top of the heart.

Electrocardiograms
we can monitor the electrical activity of the heart using an electrocardiogram (ECG). This
involves attaching a number of sensors to the skin. Some of the electrical activity generated
by the heart spreads through the tissues next to the heart and outwards to the skin. The
sensors on the skin pick up the electrical excitation created by the heart and convert this into
a trace.

The trace of a healthy person has a particular shape. It consists of a series of waves that are
labelled P, Q, R, S and T
• Wave P shows the excitation of the atria
QRS indicates the excitation of the ventricles.
• T shows diastole.

The shape of the ECG trace can sometimes indicate when part of the heart muscle is not
healthy. Such traces can be used by medical professionals to diagnose heart problems.

affinity: a strong attraction

dissociation: means releasing the oxygen from the oxyhaemoglobin
fetal haemoglobin: the type of haemoglobin usually found only in the fetus
haemoglobin: the red pigment used to transport oxygen in the blood.

Haemoglobin
Oxygen is transported in the red blood cells (erythrocytes). These cells contain the protein
haemoglobin. When haemoglobin takes up oxygen, it becomes oxyhaemoglobin.

haemoglobin + oxygen-> oxyhaemoglobin

Haemoglobin is a complex protein with four subunits. Each subunit consists of a polypeptide
chain and a haem (non-protein) group. The haem group contains a single iron ion in the
form of Fe. This iron ion can attract and hold an oxygen molecule. The haem group is said to
have a high affinity (attraction) for oxygen. As each haem group can hold one oxygen
molecule, each haemoglobin molecule can carry four oxygen molecules. It has been
calculated that there are about 280 million molecules of haemoglobin in each red blood cell
— so a red blood cell can carry over a billion oxygen molecules.

Association of oxygen and haemoglobin occurs in the lungs where pO2 is high.
Dissociation of oxygen and haemoglobin occurs in the tissues where pO2 is low.

Transport of oxygen
Oxygen is absorbed into the blood as it passes the alveoli in the lungs. Oxygen molecules
diffusing into the blood plasma enter the red blood cells. Here they become associated with
the haemoglobin. This means that the oxygen binds reversibly to the haemoglobin. This
takes the oxygen molecules out of solution and so maintains a steep concentration gradient,
allowing more oxygen to enter the blood from the lungs and diffuse into the cells.
The blood carries the oxygen from the lungs back to the heart, before travelling around the
body to supply the tissues. In the body tissues, cells need oxygen for aerobic respiration.
Therefore the oxyhaemoglobin must be able to release the oxygen. This is called
dissociation.

Haemoglobin and oxygen transport

The ability of haemoglobin to associate with and release oxygen depends on the
concentration of oxygen in the surrounding tissues. The concentration of oxygen is
measured by the relative pressure that it contributes to a mixture of gases. This is called the
partial pressure of oxygen or pO2. It is also called the oxygen tension, and is measured in
units of pressure (kPa). With a normal liquid, you might expect the concentration of oxygen
absorbed into the liquid to be directly proportional to the oxygen tension in the surrounding
air. A graph of percentage saturation plotted against oxygen tension would be a straight line
(linear). This is not the case with blood containing haemoglobin. Haemoglobin can associate
with oxygen in a way that produces an S-shaped curve. This is called the haemoglobin
dissociation curve. At low oxygen tension, the haemoglobin does not readily associate with
oxygen molecules. This is because the haem groups that attract the oxygen are in the
centre of the haemoglobin molecule. This makes it difficult for the oxygen molecule to reach
the haem and associate with it. This difficulty in combining with the first oxygen molecule
accounts for the low saturation level of haemoglobin at low oxygen tensions.

As the oxygen tension rises, the diffusion gradient into the haemoglobin molecule increases.
Eventually, one oxygen molecule enters the haemoglobin molecule and associates with one
of the haem groups. This causes a slight change in the shape of the haemoglobin molecule,
known as a conformational change. It allows more oxygen molecules to enter the
haemoglobin molecule and associate with the other haem groups relatively easily. This
accounts for the steepness of the curve as the oxygen tension rises.

As the haemoglobin approaches 100% saturation, the curve levels off, creating the S-
shaped curve. Mammalian haemoglobin is well adapted to transporting oxygen to the
tissues of a mammal. The oxygen tension found in the lungs is sufficient to produce close to
100% saturation. The oxygen tension in respiring body tissues is sufficiently low to cause
oxygen to dissociate readily from the oxyhaemoglobin.

Fetal haemoglobin
Fetal haemoglobin is slightly different from adult haemoglobin. Fetal haemoglobin has a
higher affinity for oxygen than adult haemoglobin. Therefore, the haemoglobin dissociation
curve for fetal haemoglobin is to the left of the curve for adult haemoglobin. This is because
fetal haemoglobin must be able to associate with oxygen in an environment where the
oxygen tension is low enough to make adult haemoglobin release oxygen. In the placenta,
where the oxygen tension is low, fetal haemoglobin will absorb oxygen from the surrounding
fluid. This reduces the oxygen tension even further. As a result, oxygen diffuses from the
mother's blood fluid into the placenta. This reduces the oxygen tension within the mother's
blood, which, in turn, makes the maternal haemoglobin release more oxygen (dissociation).

carbonic anhydrase: the enzyme that catalyses the combination of carbon dioxide and
water. chloride shift: the movement of chloride ions into the erythrocytes to balance the
charge as hydrogencarbonate ions leave the cell.
Bohr effect: the effect that extra carbon dioxide has on the haemoglobin, explaining the
release of more oxygen.
haemoglobinic acid: the compound formed by the buffering action of haemoglobin as it
combines with excess hydrogen ions.

The role of haemoglobin

Carbon dioxide is released from respiring tissues. It must be removed from these tissues
and transported by the blood to the lungs for excretion.
Carbon dioxide is transported in three ways:
• About 5% is dissolved directly in the plasma.
• About 10% is combined directly with haemoglobin to form a compound called
carbaminohaemoglobin.
• About 85% is transported in the form of hydrogencarbonate ions (HCO3-).

Formation of hydrogencarbonate ions

Carbon dioxide in the blood plasma diffuses into the red blood cells. Here, it combines with
water to form a weak acid called carbonic acid. This reaction is catalysed by the enzyme
carbonic anhydrase.
CO2 + H2O —> H2CO3

This carbonic acid dissociates to release hydrogen ions (H+) and hydrogencarbonate ions
H2CO3-> HCO3- + H+
The hydrogencarbonate ions diffuse out of the red blood cell into the plasma. The charge
inside the red blood cell is maintained by the movement of chloride ions (Cl-) from the
plasma into the red blood cell. This is called the chloride shift.
The hydrogen ions building up in the red blood cell could cause the contents of the red blood
cell to become very acidic. To prevent this. the hydrogen ions are taken out of solution by
associating with haemoglobin to produce haemoglobinic acid (HHb). The haemoglobin is
acting as a buffer (a compound that maintains a constant pH).

The effect of increasing carbon dioxide concentration

Blood entering respiring tissues carries oxygen as oxyhaemoglobin. The partial pressure of
oxygen in the respiring tissues is lower than that in the lungs, because oxygen has been
used in respiration. As a result, the oxyhaemoglobin begins to dissociate and releases
oxygen to the tissues. This means that the haemoglobin is available to take up the hydrogen
ions, forming haemoglobinic acid. Where the tissues are very active there is more carbon
dioxide released. This has a dramatic effect on the haemoglobin.

The Bohr effect

The Bohr effect describes the effect that an increasing concentration of carbon dioxide has
on the haemoglobin.
• Carbon dioxide enters the red blood cells forming carbonic acid, which dissociates to
release hydrogen ions.
• These hydrogen ions affect the pH of the cytoplasm, making it more acidic.
• As with any protein, changes in pH can affect the tertiary structure of the haemoglobin. The
increased acidity alters the tertiary structure of the haemoglobin and reduces the affinity of
the haemoglobin for oxygen.
• The haemoglobin is unable to hold as much oxygen, and oxygen is released from the
oxyhaemoglobin to the tissues. Where tissues (such as contracting muscles) are respiring
more, there will be more carbon dioxide. As a result, there will be more hydrogen ions
produced in the red blood cells. This makes the oxyhaemoglobin release more oxygen.

So, when more carbon dioxide is present, haemoglobin becomes less saturated with
oxygen. This is reflected in a change to the haemoglobin dissociation curve, which shifts
downwards and to the right- the Bohr shift. This Bohr effect results in more oxygen being
released where more carbon dioxide is produced in respiration. This is just what the muscles
need for aerobic respiration to continue.

Exam Prep

Transport in animals
 Oxygen is not released by erythrocytes until they reach the capillaries as arteries have a thick wall whereas the
capillary wall is only one cell thick, so the diffusion distance would be too large for the artery, but a is short enough
diffusion distance to occur through the capillary wall.
 Advantages of erythrocytes having no nuclei or membrane-bound organelles include that this leaves more space for
haemoglobin and thus oxygen, and also allows them to squeeze through capillaries easily
 Disadvantages of erythrocytes having no nuclei or other membrane-bound organelles include iti gives them a limited
life span, they can’t reproduce/divide /undergo mitosis, can’t do any protein synthesis / repair, and no mitochondria
for respiration
 Erythrocytes don’t use the oxygen they carry as oxygen is bound to haemoglobin while being transported so cannot
be used, and also they lack mitochondria, therefore can’t use the oxygen for aerobic respiration. Also erythrocytes
are moved by mass flow so they don’t need ATP produced from respiration to move and need less ATP for metabolic
processes due to lack of organelles
 Blood offloads more oxygen to actively respiring tissues than resting tissues as in actively respiring tissues there are
higher levels of carbon dioxide, which lower the affinity of haemoglobin for oxygen, resulting in dissociation of
carbonic acid and increase of H+, leading to the release of oxygen, so more oxygen released at same pO2
 To compare frog and mammalian circulation state how they are similar, how they are different, and then comment on
which is more effective and why
 Both frogs and mammals have double circulations, though blood from mammalian heart transported separately to
lungs and body and oxygenated and deoxygenated blood never mix, whereas blood from frog heart transported to
lungs and body together so the blood going to the body in the frog is partially oxygenated and oxygenated blood only
separate when returning from lungs. In frogs the spiral valve partly separates oxygenated and deoxygenated blood
 In terms of effectiveness both can be considered to be effective frog could be considered to be less effective as frog
has less oxygen available for the body cells, but the circulation is effective enough for the frog’s needs as the frog
has lower metabolic rate and as the frog maintains body temperature by other means, and as the frog heart may beat
faster to compensate. However frog circulation may limit its size. Mammalian system more effective as mammalian
body cells get maximum available O2, though this is necessary as the mammal has a higher metabolic rate and uses
metabolism to maintain body temperature
 Electrical activity in the heart is controlled by a wave of depolarisation from the AVN moving along the Purkyne fibres
to cause the contraction of the ventricles.
 In a dissection it is better to use the heart that has blood vessels present
 Diastole follows systole in the mammalian heart because cardiac muscle takes a short time to repolarise after being
stimulated
 In a double circulatory system blood flows twice through the heart for one heart beat.
 High blood pressure causes cell death as increased blood pressure causes small blood vessels such as capillaries to
burst / break, causing a localised build up of pressure leading to cell death. In this place blood cannot circulate to
supply oxygen, so cells cannot respire, leading to cell death
 The two circuits are the pulmonary and systemic systems
 A toxin that reduces speed of conduction down the purkyne fibres slows down heart rate as slows transmission of
impulse from AVN to ventricles, slows ventricular systole / contraction, longer delay before ventricular systole
/contraction begins, and increases time the heart is in diastole / relaxation
 The left ventricle is much larger than the right ventricle and the right atrium as the wall of cardiac muscle is thicker,
and it provides higher pressure to pump blood around body as it needs to pump blood a longer distance so there is
greater resistance than for the shorter distance that the right ventricle or right atrium has to pump blood
 The left ventricle is much thicker than the left atrium as more muscle to create more force as it needs to create a
higher pressure to push blood against greater resistance because the left ventricle pumps blood further as it supplies
systemic circulation
 The atrioventricular valves close when ventricular systole occurs, as this raises ventricular pressure such that it is
higher than atrial pressure, and the pressure generated by ventricular contraction pushes the valves shut, and
chordae tendinae prevent inversion
 In a single circulatory system blood passes through the heart once per circuit of the body
 In a closed circulatory system the blood is maintained inside vessels
 In a heartbeat, the sinoatrial node initiates the excitation. The wave of excitation spreads over atrial walls causing
them to contract in a synchronised manner (atrial systole). Non-conducting tissue forces the wave of excitation to
pass through the atrioventricular node, so the excitation spreads down the septum through the Bundle of his and then
through the Purkyne fibres in the walls of the ventricles, causing the ventricles to contract (ventricular systole) from
apex of the upwards.
 Cardiac muscle is found in the walls of the heart chamber
 Ventricular and atrial systole cause pressure inside the heart chambers
 Arteries differ to veins as they have no valves, their endothelium is more folded, they have thicker muscle tissue and
thicker elastic tissue, and they also have a thicker collagen layer.
 Histamine stimulates smooth muscle contraction by attaching to complementary receptors on the cell surface
membrane of muscle cells, triggering a response inside the cells, it also makes capillary walls more permeable
 The permeability caused by the histamine leads to more tissue fluid formed, so increase in pressure in the tissue, so
swelling / inflammation / oedema, and more larger molecules like white blood cells pass into the tissue fluid
 Hydrostatic pressure in the arteries is generated by contraction of the ventricle muscle wall in the heart
 Hydrostatic pressure drops as blood moves away from the heart as there are more vessels the further you get from
the heart, so the vessels have a larger total lumen/cross-sectional area, also there is reduced resistance to blood
flow, the arteries stretch/expand, and fluid is lost as plasma from the capillaries
 At the arterial end of the capillary plasma moves out of the capillary and forms tissue fluid. Plasma proteins remain in
capillary as they are too large to pass through capillary wall. The fluid moves down pressure gradient as hydrostatic
pressure is greater than water potential here
 Advantages of keeping blood in vessels include maintaining a higher blood pressure to increase rate of flow/delivery,
and also allows flow to be directed/diverted
 The artery is adapted to withstand pressure as its wall is thick, with a thick layer of collagen. The collagen provides
strength. The endothelium is also folded so it can stretch
 The artery is adapted to maintain pressure as the thick layer of elastic tissue allows the artery to recoil back to its
original size. There is also a thick layer of smooth muscle to allow the artery lumen to constrict
 Pressure fluctuates in the aorta due to systole increasing pressure via contraction of the left ventricle wall and
diastole decreasing pressure
 The number of pressure fluctuations per minute is the pulse/heart rate
 It is important that blood pressure changes as blood flows from the aorta to the capillaries as the capillary wall is thin,
so high pressure would damage the capillary wall, so lower pressure reduces the risk of tissue fluid build up
 Oxygen enters red blood cells by diffusion
 There is a delay between the excitation of the atria and the excitation of the ventricles to allow time for the atria to
fully contract and empty of blood and to allow time for the ventricles to fill with blood so they do no contract too early
 The excitation wave travels down the septum to the apex of the heart so that ventricular contraction starts at the apex
of the heart to push blood upwards into the arteries and allow for complete emptying of the ventricles
 People with sickle cell anaemia benefit from drugs that cause them to produce fetal haemoglobin as fetal
haemoglobin may not crystallise much at low pO2, so these red blood cells do not change shape and so cause
symptoms of sickle cell anaemia. Also fetal haemoglobin can pick up more oxygen at lower pO2 than sickle
haemoglobin, so more oxygen is transported around the body
 Substances dissolved in blood that are dissolved in the blood plasma are able to enter the tissue fluid via diffusion
from high concentration to low concentration down the concentration gradient as hydrostatic pressure in the capillary
is higher than in the tissue fluid, and the capillary walls are leaky so fluid is forced out of the capillary down the
pressure gradient. As the fluid moves out the glucose, oxygen, and other small molecules, leave with it
 Erythrocytes aren’t found in tissue fluid because the gaps between endothelial cells are too small for the erythrocytes
to fit through, and the erythrocytes can’t change shape much so can’t bend to fit through the fenestrations between
the endothelial cells
 Haemoglobin has a high affinity for oxygen, so oxygen binds to haemoglobin in the alveoli in the lungs where the pO2
is high, forming oxyhaemoglobin. Oxygen is then released in tissues where respiration is occuring and so it is needed
as the pO2 there is low
 Hydrogencarbonate ions are produced in the erythrocytes as carbon dioxide diffuses into the erythrocytes and reacts
with water with help from the enzyme carbonic anhydrase, forming carbonic acid. The carbonic acid then dissociates
to form hydrogencarbonate ions and hydrogen ions
 The Bohr effect occurs as carbon dioxide reduces the affinity of haemoglobin for oxygen. The protons from the
dissociation of carbonic acid interact with haemoglobin to form haemoglobinic acid as this prevents the fall of pH in
the cells in the surroundings, so provides a buffering effect. The H+ ions alter the structure of haemoglobin such that
more oxygen released where it is needed more (i.e: respiring tissues) as carbon dioxide concentration is high in
these areas. CO2 then binds to haemoglobin forming carbaminohaemoglobin
 If the ventricles contract twice for every atrial contraction in the heart, then less blood leaves heart for each
ventricular contraction as the ventricles do not have time to fill before contracting
 The hole in the septum between heart chambers is open in a fetus before birth as the lungs are not functioning as
they are not filled with air, so blood is not oxygenated in the lungs, and instead is oxygenated in the placenta,
therefore pulmonary circuit / lungs are bypassed
 Fetal haemoglobin contains gamma subunits to give it a higher affinity for oxygen than adult haemoglobin
 There are two blood vessels carrying blood away from the heart as the mammalian body has a double circulatory
system. One circulation goes to the lungs (pulmonary circuit), and one goes to the body (systemic circuit)
 Weak irregular heartbeat may result in fatigue as it results in slow blood flow and lower blood pressure, so less
oxygen reaches the cells for oxidative phosphorylation in aerobic respiration, so less glucose reaches the cells for
respiration, so less ATP produced. There is then also increased acidity as CO2 / lactate builds up which affects the
enzymes involved in respiratory metabolism