Capillary Function and Blood Gases
· Blood consists of red blood cells (erythrocytes), white
blood cells (lymphocytes) and platelets, suspended in the liquid blood plasma.
· Red cells contain
haemoglobin and are involved in the transport of oxygen and carbon dioxide by
the blood.
· White cells are a major part
of the immune system.
· Platelets are cell fragments
involved in clotting of the blood when blood vessels are damaged.
· Blood plasma is mainly water.
· It contains many substances
in solution, including;
· ions - sodium, potassium,
chloride, bicarbonate and others,
· t
· hormones - insulin, glucagon
and many others,
· urea,
· proteins.
· The proteins are of various types, including;
· enzymes,
· antibodies (to fight
infections),
· albumins,
· globulins (to carry lipids),
· fibrinogen.
· When blood clots, the fibrinogen changes into insoluble
threads of fibrin.
· These threads form a mesh
over a break in the wall of a blood vessel.
· Platelets get caught in the
mesh and a clot forms.
· Blood serum is what remains when the fibrin/fibrinogen is
removed from blood plasma.
· Tissue fluid is the liquid which is found
between/surrounding all the cells of the body.
· It is formed from components
of the blood plasma which leave the blood as it passes through the capillaries.
· Most of the large components
of the blood can not cross the capillary wall, so tissue fluid does not contain
blood cells (except for certain white cells), platelets, or large plasma
proteins.
· Water, mineral ions, glucose
and amino acids can cross the wall and are found in similar concentrations to
those in blood plasma.
· Tissue fluid has a lower
concentration of oxygen and a higher concentration of carbon dioxide than
plasma.
· Lymph is formed when surplus tissue fluid enters lymph
capillaries.
· These capillaries are
dead-end vessels which merge to form larger lymph vessels.
· The lymph is eventually
returned to the blood, to form part of the plasma again.
· Lymph is very similar in
composition to tissue fluid but may contain fats (absorbed in digestion), more
protein and white blood cells (in the lymph nodes).
The exchange of substances between the blood
and tissue fluid across the wall of the capillary.
· At the arterial end of a capillary, the blood pressure is relatively high.
· This hydrostatic pressure forces water through the wall of the capillary
into, and becoming part of, the tissue fluid.
· Glucose and mineral ions
leave by diffusion.
· Oxygen diffuses out into the
tissue fluid along a concentration gradient - since the cells are constantly
using up oxygen in respiration.
· Carbon dioxide and urea
diffuse into the plasma along concentration gradients.
· Blood cells and plasma
proteins are too large to get through the capillary wall.
· The loss of water lowers the
volume of the blood and this, together with friction, reduces the blood
pressure as the blood flows towards the venous end of the capillary.
· The loss of water also
increases the concentration of the plasma proteins, making the water potential of the blood more
negative.
· This lower blood pressure
and water potential allows water to flow back into the blood from the tissue
fluid, along a water potential gradient
by osmosis. You may have been taught this as Starling’s Hypothesis.
· The water potential gradient
is not large enough to reabsorb all of the water that leaves the capillary.
· This means that the volume
of the tissue fluid would tend to increase (and blood volume decrease).
· The surplus tissue fluid is
drained away by the lymphatic system and returned to the blood.
You
should understand how oxygen and carbon dioxide are loaded, carried and
unloaded by the blood. You should understand the buffering effect of
haemoglobin.
· Oxygen. The vast majority of the oxygen in the blood is
carried in red blood cells (erythrocytes).
· It is chemically bound to haemoglobin inside the cells.
· Each haemoglobin molecule
can hold up to four oxygen molecules.
· Haemoglobin is a protein with specific receptor/binding sites on its surface for oxygen.
· The oxygen fits into the receptor
sites because its shape fits that of
the receptor site.
Note. Haemoglobin is not an
enzyme but many of its characteristics are similar to an enzyme’s.
· Oxyhaemoglobin is formed when oxygen binds to haemoglobin.
Haemoglobin + Oxygen Û
Oxyhaemoglobin
· The binding of oxygen is reversible,
· depending upon the concentration of oxygen and the strength of the
affinity/attraction of haemoglobin for oxygen.
· Blood entering the lung capillaries has little oxygen bound to its haemoglobin.
· The capillaries surround the
alveoli of the lungs, which contain
a high concentration of oxygen.
· This creates a concentration gradient for the diffusion of oxygen.
· Oxygen dissolves in the
liquid lining the inside of the alveoli and then diffuses through the wall of the alveoli and capillary, into the
blood plasma.
· It then diffuses into the red cells and is bound to haemoglobin.
· This takes it out of
solution, effectively keeping the concentration of dissolved oxygen low in the
blood and maintaining the concentration
gradient.
· As the haemoglobin becomes
saturated with oxygen, it is carried away from the lungs as the blood flows and
replaced with more oxygen poor haemoglobin.
· This also maintains the concentration gradient for
oxygen diffusion.
· The haemoglobin in the lung capillaries has a high affinity for oxygen,
due to the excretion of carbon dioxide by the lungs and the resulting changes in
blood chemistry (see carbon dioxide transport later in this section).
· Respiration takes place all the time in the tissues of the
body.
· This means that the cells
are continuously using oxygen.
· As a result, the
concentration of oxygen in the tissues is lower than in the blood.
· A concentration gradient exists for the diffusion of oxygen from the
red cells, into the blood plasma, the tissue fluid and the cells of the
tissues.
· The lowering of the oxygen
concentration in the red cells causes the reversal
of the binding of oxygen to haemoglobin.
· Respiration in the tissues
also increases the concentration of carbon
dioxide.
· This produces changes in
blood chemistry which lower the affinity
of haemoglobin for oxygen, causing more oxygen to be released, more easily.
· Carbon dioxide. Carbon dioxide is carried in the blood in
three ways.
· Dissolved carbon dioxide - about 10% is carried as
dissolved carbon dioxide molecules in the blood plasma.
· Carbamino-haemoglobin - about 20% is carried
attached to an amino acid in the haemoglobin molecule.
· Hydrogencarbonate ions - about 70% is carried as
these ions in solution in the blood plasma.
· Respiration in the tissues produces a high concentration
of carbon dioxide.
· This creates a concentration gradient for the diffusion
of carbon dioxide from the tissues and into the blood plasma.
· Most of the carbon dioxide
then diffuses into the red cells, because of a low concentration of carbon dioxide
in these cells.
· The concentration is low,
because red cells contain an enzyme, carbonic
anhydrase, which speeds up the reaction of carbon dioxide with water to
produce carbonic acid (effectively ‘using-up’ the carbon dioxide).
· The carbonic acid then
dissociates to produce hydrogencarbonate
ions and H+ ions.

· The reaction is reversible,
depending upon the concentration of carbon dioxide, and this is important in the
lungs (see later).
· The hydogencarbonate ions diffuse out of the red cells into the plasma,
along a concentration gradient.
· This outflow of negative
ions would leave the red cells with an electrical charge imbalance.
· To keep the charge balance,
negative chloride ions (from sodium chloride) diffuse into the red cells.
· This is the chloride shift.
· The H+ from the
carbonic acid could be a problem, since they tend to lower the pH of (make more
acidic) the red cells and the blood.
· Haemoglobin helps to prevent
the pH change by binding to H+.
· This is haemoglobin acting as a buffer.
· Binding to H+
lowers the affinity of haemoglobin for oxygen, making it easier for oxygen to
be supplied to the tissues.
· The greater the rate of
respiration in the tissues, the greater the concentration of carbon dioxide,
the more H+ is produced and the more easily oxygen is released to
the tissues that need it.
· This increase in
release/dissociation of oxygen from oxyhaemoglobin caused by increases in
carbon dioxide concentration is the Bohr
effect.
· When the blood reaches the lung
capillaries, carbon dioxide diffuses
out of the blood plasma into the alveoli, along a concentration gradient.
· This lowers the carbon
dioxide concentration in the plasma and causes the reactions involving carbonic anhydrase to go into reverse.
· As a result,
hydrogencarbonate ions are converted back into carbon dioxide and excreted into
the alveoli.
· The other events, involving
H+ and chloride ions are also reversed.
In exam
questions, the formation of hydrogencarbonate ions might be shown on a diagram
of a red cell.
Note that all of the reactions involved in these events
are synoptic with sections 1.3, 1.4 and 1.5 of By01.
You should be able to interpret
oxygen-haemoglobin dissociation curves.
·
The graph shows a dissociation curve for human haemoglobin.
·
Many organisms have haemoglobin but different to human haemoglobin, each
with its own dissociation curve. (This is discussed later.)

·
The curve is sigmoid/S-shaped.
·
The partial pressure of oxygen can be thought of as the ‘concentration’
of oxygen.
·
In the lungs and major arteries the % saturation is about 97%.
·
Because the graph levels off
at the top, it means that haemoglobin becomes fully saturated
with oxygen over a wide range of atmospheric partial pressures of oxygen.
·
This is why you can climb quite high mountains without the need for an
oxygen mask!
·
When the body is at rest, the tissues remove oxygen for respiration and
leave the haemoglobin about 75% saturated with oxygen.
·
This corresponds to an oxygen partial pressure of just over 5kPa.
·
During exercise, the tissues use far more oxygen, lowering the partial
pressure to about 4kPa.
·
From the graph, you can see that this small fall in partial pressure
causes a large fall in the saturation of haemoglobin from 75% to about 58%.
·
This is the significance of
the steep part of the dissociation curve: a small fall in oxygen in the tissues
causes a lot of oxygen to dissociate from haemoglobin.
·
A rise in the rate of
respiration causes much more oxygen to be released into the tissues, to
maintain the high rate of respiration.
·
A high rate of respiration also produces more carbon dioxide.
·
This is converted into carbonic acid in the red cells.
·
As discussed earlier in this section, this lowers the affinity of
haemoglobin for oxygen - the Bohr
Effect.


· The graph shows the effect
of an increase in the acidity of the
blood on the dissociation curve for haemoglobin.
· The curve has moved to the right.
· The more carbon dioxide
produced/ the greater the fall in the pH, the greater the shift to the right.
· As you can see from the
graph, a fall in pH of 0.2 causes 18% extra
oxygen to dissociate (it falls from
63% to 45%) from haemoglobin at a partial pressure of oxygen of 4kPa in the
tissues.
· Again, this extra oxygen would help to maintain the
high rate of respiration producing the carbon dioxide which lowered the pH.
· You might be given data from
different species of organism living in different environments, with different
amounts of oxygen available.
· For example, fish live in
water and this contains much less oxygen than air.
· The fish have haemoglobin
which becomes fully saturated at much lower partial pressures of oxygen than
the haemoglobin of air-breathing mammals.
· Worms that live in mud flats
(e.g. Lugworms) have even less oxygen available in their environment than fish.
· They have evolved
haemoglobin which is fully saturated at a very low partial pressure of oxygen;
much lower than for fish.
· So, the dissociation curve
for fish haemoglobin would be to the left of that for human haemoglobin.
· The curve for the worm would
be to the left of that for the fish.
· You might be given data on
another ‘special case’, the human foetus.
· The human foetus has slightly
different haemoglobin to that of humans once they have been born.
· Foetal haemoglobin has a dissociation curve to the left of
‘normal’ human haemoglobin.
· This means that it has a higher affinity for oxygen than the
mother’s blood.
· This allows the foetus to
take oxygen from the mother’s blood, across the placenta.
Common
Mistakes
Not knowing in sufficient detail how carbon dioxide is
carried in the blood.
Not appreciating that the reactions involved in
carrying oxygen and carbon dioxide are reversible. The conditions in the
alveoli of the lungs and the tissues are the opposite of each other: high
oxygen concentration/partial pressure in the alveoli and low in the respiring
tissues, high carbon dioxide concentration/partial pressure in the tissues and
low in the alveoli. These conditions are linked to diffusion gradients created
by respiration in the tissues (using up oxygen and producing carbon dioxide)
and ventilation of the lungs (removing carbon dioxide and bringing in oxygen).
Not appreciating that the shape of the haemoglobin
dissociation curve and changes such as the Bohr effect are linked to providing
oxygen to rapidly respiring tissues.
Practice
Questions
|
1. |
Explain how carbon dioxide produced in respiring
tissues is loaded and carried in the blood and finally unloaded in the lungs.
(6 marks) (15 lines were allowed for the answer.) |
Northern
Examinations and Assessment Board June‘96. Question 8. [Part]
|
2. |
The graph shows the oxygen-haemoglobin dissociation
curves for a mouse and some other mammals. |
Key: 1 Elephant 2 Horse 3 Cat 4 Mouse


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a) |
Explain how the haemoglobin of the mouse is able to
take up oxygen in the lungs and unload it in the tissues. (5 marks) (10 lines were allowed for the answer.) |
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b) |
i) |
Describe the relationship between the position of the
oxygen-haemoglobin dissociation curve and body size for the species shown in
the graph. (1 mark) |
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ii) |
Explain the advantage to a mouse of having an
oxygen-haemoglobin curve of this shape and position. (3 marks) (7 lines were allowed for the answer.) |
Northern
Examinations and Assessment Board June‘97. Question 9.Part.
|
3. |
Nephrosis is a kidney condition in which damage to
the glomeruli results in large quantities of protein passing into the glomerular
filtrate. This protein finally appears in the urine. |
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a) |
Suggest why this protein is not reabsorbed into the
blood in the proximal convoluted tubule of the nephron. (1 mark) |
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b) |
As a result of nephrosis large amounts of tissue
fluid accumulate in the body, especially in the ankles and feet. |
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i) |
Explain why the loss of protein from the blood
results in the accumulation of tissue fluid. (2 marks) |
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ii) |
Suggest why this fluid accumulates especially in the
ankles and the feet. (2 marks) |
Northern
Examinations and Assessment Board June‘95. Question 6. [Part]
|
4. |
The graph shows oxygen-dissociation curves for haemoglobin
as blood passes through capillaries in the lungs and in the skeletal muscles
of an athlete. |


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Explain how features of the oxygen-dissociation
curves for haemoglobin in the lungs and in the skeletal muscles benefit an
athlete. (7 marks) (18 lines were allowed for the answer.) |
Northern
Examinations and Assessment Board March‘98. Question 8.Part.
Model
Answers
|
1. |
Carbon dioxide diffuses into the blood (in
solution); where some is carried in solution as carbon dioxide; most moves into red blood cells along a
concentration gradient; there is carbonic anhydrase (in the red cells); which converts carbon dioxide into carbonic
acid/hydrogencarbonate ions; the hydrogencarbonate ions diffuse out into the
blood plasma; this is the major form in which carbon dioxide is
carried; some is carried attached/bound to haemoglobin
molecules; the movement of hydrogencarbonate out of the red
cells is balanced by chloride ions moving in/the chloride shift; in the lungs the reactions involving
hydrogencarbonate are reversed (because of the low partial pressure of carbon dioxide in the
alveoli); carbon dioxide diffuses out into the alveoli; (any 6 points) |
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2. |
a) |
Oxygen diffuses into the blood (from the alveoli); the high partial pressure of oxygen in the lungs
means that haemoglobin becomes saturated fully loaded with oxygen; each molecule of haemoglobin can carry four oxygen
molecules; forming oxyhaemoglobin; in the tissues, the oxygen partial pressure is lower
(due to respiration) and oxygen is released/unloaded from oxyhaemoglobin; the higher carbon dioxide levels (from respiration)
in the tissues shift the dissociation curve for haemoglobin to the right/Bohr shift; this causes more oxygen to be unloaded from
haemoglobin for respiration; the higher carbon dioxide levels also mean more
carbonic acid production, producing H+ ions which also favours the release of oxygen
from haemoglobin (as a buffer); (any 5 points) |
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b) |
i) |
The smaller the animal, the further the curve is to
the right; |
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ii) |
Small animals lose more heat; which needs a higher rate of respiration, to replace
the lost heat; this needs more oxygen; their haemoglobin releases its oxygen more easily; (any 3 points) |
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3. |
a) |
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The protein molecules are too large to pass through the
membranes of the cells of the tubule; |
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b) |
i) |
The loss of protein makes the water potential of the
blood plasma less negative; this means that less tissue fluid is reabsorbed into
the blood in the capillaries; |
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ii) |
There are not many lymph vessels in the ankles/feet; and so the tissues do not drain; |
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OR |
There are few muscles in the ankles/feet; so not much squeezing of tissues to drain them; |
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4. |
Haemoglobin in red cells returning to the lungs has
little oxygen, and thus a high affinity for oxygen; it becomes saturated with oxygen in the lungs; and this happens over quite a wide range of (lung)
oxygen concentrations (as seen by the flat top to the curve); in the muscle capillaries there is a high carbon
dioxide concentration/low pH (due to a high rate of respiration); which produces a Bohr effect/shifts the curve to the
right; this reflects a lower affinity of haemoglobin for
oxygen; so, oxyhaemoglobin dissociates/gives up oxygen
easily; which gives more oxygen for respiration in the
muscle cells; (any 7 points) |