Gaseous exchange

Transport of oxygen
The red blood cells contain haemoglobin, the molecule that transports oxygen. Typically, the human body requires
between 250 cm3 (at rest) and 1000 cm3 (in strenuous exercise) of oxygen per minute. At human
(37 oC) body
temperature, about 0.45 cm3 of oxygen can be dissolved in 100 cm3 of blood plasma, but the total oxygen transporting
facility of blood is about 20 cm3 per 100 cm3. Thus some 98 per cent of oxygen is transported by haemoglobin.
Haemoglobin and oxygen transport
Haemoglobin is a compact molecule made up of four interlocking sub-units. The sub-units consist of four polypeptide
chains called globin (two -chains and two -chains), each attached to a haem group. Haemoglobin therefore has a
quaternary structure. The haem groups comprise a porphyrin ring containing an atom of iron (II) at the centre. The haem
groups in haemoglobin are responsible for its red colour, and are the site of oxygen transport. An iron atom combines with
a molecule of oxygen, but without oxidation of the iron (II). Thus, each haemoglobin molecule can carry four molecules
of O2. Like all respiratory pigments, haemoglobin must bind oxygen reversibly, loading oxygen in the lungs and unloading
it in other parts of the body. Loading and unloading depends on cooperation among the sub-units of the haemoglobin
molecule. The binding of oxygen to one sub-unit induces the remaining sub-units to change their shape slightly so that
their affinity for oxygen increases. And when one sub-unit unloads its oxygen, the other three quickly follow suit as a
conformational change lowers their affinity for oxygen.
The structure of haemoglobin
Haemoglobin is a conjugated protein
The protein part (called globin) consists of four polypeptide chains. These chains are of two types called alpha
and beta. They are about the same length (about 140 amino acids) but have slightly different compositions.
Each chain is combined with a non-protein prosthetic group called haem. Haem consists of an atom of iron
enclosed in a ring structure (porphyrin ring)
Each haem group can combine with one molecule of oxygen. This process is called oxygenation. (It is not the
same as oxidation, because the iron does not lose any electrons and is not chemically oxidised). Each molecule of
haemoglobin can therefore combine with a maximum of four molecules of oxygen.
Oxygen tension (partial pressure)
In a mixture of gases, each component gas exerts a pressure (its partial pressure or tension) in proportion to its
molar percentage in the mixture.
The partial pressure of a gas is the pressure contributed by one gas to the total pressure of a mixture of gases.
The partial pressure of a gas is the pressure exerted by the particular gas in a mixture of gases.
Partial pressures of the components of air
Component gases Percentage composition Partial pressure/kPa Partial pressure/mm Hg
Oxygen
21
21.2
160
Carbon dioxide
0.04
0.04
0.3
Nitrogen
79
80.0
Partial pressure of the components of alveolar air
Components gases Percentage composition Partial pressure/kPa Partial pressure/mm Hg
Oxygen
13.8
14.0
104
Carbon dioxide
5.5
5.6
40
Nitrogen
80.7
Partial pressures of the components in tissue cells
Component gases Percentage composition Partial pressure/kPa Partial pressure/mm Hg
Oxygen
< 5.6
< 40
Carbon dioxide
> 6.3
> 45
Oxygen transport
The Po2 of the air within alveoli at sea level is approximately 105 mm Hg (14 kPa), which is less than the Po 2 of the
atmosphere because of the mixing of freshly inspired air with old air in the anatomical dead space of the respiratory
system. The Po2 of the blood leaving the alveoli is slightly less than this, about 100 mmHg, because the blood plasma is
not completely saturated with oxygen due to slight inefficiencies in lung function. At a blood Po 2 of 100 mm Hg,

approximately 97% of the haemoglobin within red blood cells is in the form of oxyhaemoglobin indicated as a percent
oxyhaemoglobin saturation of 97%.
As the blood travels through the systemic blood capillaries, oxygen leaves the blood and diffuses into the tissues.
Consequently, the blood that leaves the tissue in the veins has a Po 2 that is decreases (in a resting person) to about 40 mm
Hg (5.6 kPa). At this lower Po2, the percent saturation of the haemoglobin is only 75%. A graphic representation of these
changes is called an oxygen (oxyhaemoglobin) dissociation curve. In a person at rest, therefore, 22% (97% minus 75%) of
the oxyhaemoglobin has releases its oxygen to the tissues. Put another way, roughly one-fifth of the oxygen is unloaded in
the tissues, leaving four-fifths of the oxygen in the blood as a reserve.
This large reserve of oxygen serves two important functions. First, it enables the blood to supply the bodys
oxygen needs during exercise as well as at rest. During exercise, the muscles accelerated metabolism uses more oxygen
from the capillary blood and thus decreases the venous blood Po 2. For example, the Po2 of the venous blood could drop to
20 mm Hg; in this case, the percent saturation of haemoglobin will be only 35%. Since arterial blood still contains 97%
oxyhaemoglobin (ventilation increases proportionately with exercise), the amount of oxygen unloaded is now 62% (97%
minus 35%), instead of the 22% at rest. The second function of the oxygen reserve is that it ensures that the blood
contains enough oxygen to maintain life for four to five minutes if breathing is interrupted or if the heart stops pumping.
The CO2 produced by metabolising tissues as a product of aerobic respiration combines with H 2O to form
carbonic acid (H2CO3), lowering the pH of the blood. This reaction occurs primarily inside red blood cells, where the
lowered pH reduces haemoglobins affinity for oxygen and thus causes it to release oxygen more readily. The effect of pH
on haemoglobins affinity for oxygen is known as the Bohr effect and is shown graphically by a shift of the oxygen
(oxyhaemoglobin) dissociation curve to the right. Since skeletal muscles produce carbon dioxide more rapidly during
exercise, the blood unloads a higher percentage of the oxygen it carries during exercise as a result of the Bohr effect
(shift).
The Bohr effect
An increase in carbon dioxide concentration will reduce the affinity of haemoglobin for oxygen, moving (shifting) the
oxygen dissociation curve of haemoglobin to the right. This causes the release of more oxygen of the respiring tissues.
Summary
Haemoglobin has a high affinity for oxygen where the partial pressure of oxygen is high. This means that it takes up
oxygen easily where it needs to, in the lings. However in the respiring tissues where the partial pressure is low, it has a
low affinity and it releases the oxygen rapidly to the cells which require the oxygen for respiration.
Oxygen dissociation curve
The degree to which the haemoglobin is saturated at different oxygen partial pressures can be measured. The oxygen
dissociation curve is S-shaped, or sigmoid.
Over the range of oxygen partial pressures (Po2) where the dissociation curve has a steep slope, even a slight change in Po 2
causes haemoglobin to load or unload a substantial amount of oxygen. Notice that the steep part of the curve corresponds
to the range of oxygen partial pressures found in body tissues.
Remember: The partial pressure of oxygen which produces 95% saturation is known as the loading tension, whereas that
producing 50% saturation is known as unloading tension.
Oxygen dissociation curves for the haemoglobin of three mammals
Llama (displaced to th left of human haemoglobin)
An animal living at high altitude where the reduced pressure of the atmosphere makes it difficult to load
haemoglobin. The llamas haemoglobin therefore has a high affinity for oxygen to compensate for this.
Mouse (displaced to the right of human haemoglobin)
Being small it has large surface area: volume ratio and therefore loses heat rapidly. To compensate for this it has a
high metabolic rate to help generate heat. To keep tissues supplied with oxygen the haemoglobin of the mouse has
a lower affinity for oxygen and therefore unloads it quickly to the tissues. The reduced affinity for oxygen does
not adversely affect the loading of haemoglobin as the oxygen tension air is 21 kPa more than enough to fully
saturate the haemoglobin.
Oxygen dissociation curves for the haemoglobin of three animals from different groups
Arenicola (annelid) (displaced to the left of human haemoglobin)
Lives in muddy waterlogged burrows where the oxygen tension is very low. If it is to obtain oxygen at all its
haemoglobin must have a very high affinity for it.
Pigeon (bird) (displaced to the right of human haemoglobin)
As flight demands much energy, birds have high metabolic rates. To supply oxygen to the tissues rapidly their
haemoglobin must release it readily. With an oxygen tension of 21 kPa the air still saturates the haemoglobin with
oxygen despite its lowered affinity for it.

Comparison of the oxygen dissociation curves of adult and foetal haemoglobin

Foetal haemoglobin has a greater affinity for oxygen than adult haemoglobin to enable it to obtain oxygen from
the mothers haemoglobin in the placenta.
Comparison of the dissociation curves of human haemoglobin and myoglobin
Myoglobin consists of a single polypeptide chain and a single haem group, rather than the four found in
haemoglobin. Dissociation curve of myoglobin is displaced well to the left of haemoglobin. Myoglobin has a
higher affinity for oxygen than haemoglobin. Myoglobin occurs in the muscles of all vertebrates. It acts as a store
of oxygen in resting muscle, only releasing it when supplied of oxyhaemoglobin have been exhausted, i.e. during
periods of strenuous exertion when the partial pressure of oxygen falls very low (below 20 mm Hg).
Transport of carbon dioxide
In solution (5%) (7%)
Most of the carbon dioxide carried in this way is transported in physical solution.
Combined with protein (10 20%) (23%)
Carbon dioxide combines with the amino group (NH2) at the end of each polypeptide chains of haemoglobin to form a
neutral carbamino-haemoglobin compound. The amount of carbon dioxide that is able to combine with haemoglobin
depends on the amount of oxygen already being carried by the haemoglobin. The less oxygen being carried by the
haemoglobin molecule, the more carbon dioxide that can be carried in this way.
As hydrogen carbonate (85%) (70%)
Carbon dioxide produced by the tissues diffuses into the bloodstream and passes into the red blood cells where it
combines with water to form carbonic acid. This process in catalysed by the enzyme carbonic anhydrase found in the
red blood cells. It is an extremely efficient enzyme. Some of the carbonic acid then dissociates into hydrogen and
hydrogen carbonate ions.
The hydrogen ions produced combine with haemoglobin which loses its oxygen (At low oxygen partial pressure
haemoglobin has a higher affinity for hydrogen ions than oxygen). The oxygen so released diffuses out of the red
blood cells, through the capillary wall and tissue fluid into a respiring tissue cell. The deoxygenated haemoglobin
accepts hydrogen ions from carbonic acid forming haemoglobinic acid (H.Hb). By accepting hydrogen ions,
haemoglobin acts as a buffer molecule and so enables large quantities of carbonic acid to be carried to the lung
without any major change in the blood pH.
The majority of hydrogen carbonate ions formed within the red blood cells diffuse out into the plasma along a
concentration gradient and combine with sodium in the plasma to form sodium hydrogen carbonate. The loss of
negatively charged hydrogen carbonate ions from the red blood cells leaves them with a more positive charge. This is
balanced by chloride ions diffusing into the red blood cells from the plasma. This phenomenon is called the chloride
shift.
In the lungs the concentration of oxygen is high due to ventilation and oxygen diffuses from the lungs into the red
blood cells. Because of the high partial pressure of oxygen, the haemoglobin has a higher affinity for oxygen, so it
(HHb) readily combines with oxygen to give oxyhaemoglobin and this displaces the hydrogen ions. These displaced
hydrogen ions combine with hydrogen carbonate ions to give carbonic acid. Catalysed by carbonic anhydrase, the
carbonic acid dissociates to produce water and carbon dioxide. The carbon dioxide diffuses from the red blood cells to
the lungs and is removed from the body. As the hydrogen carbonate ions are used up in the red cells, more diffuse in
from the plasma.
Exercises
1
(a)
Temperature also affects the dissociation of haemoglobin. Bearing in mind the effect of carbon dioxide,
suggest what the effect of a rise in temperature might be, and explain the physiological advantage of the change.
(b)
The oxygen dissociation curve is not the same for all animals. For example, compared with humans, the
curve for small mammals is displaced to the right. Suggest why this is the case.
2
The oxygen dissociation curve of the fetus is to the left of that of its mother. Suggest why this is so.
3ich lives in the High Andes at an altitude of about 5000m above sea level, is located to the left of most other mammals.
Suggest whys this is so.

Answers
1
(a)
Increased metabolic activity increases the temperature in a part of the body. This produces a reduction in
the affinity for oxygen for haemoglobin and an increased dissociation of oxygen. Thus the dissociation curve is shifted to
the right. This is physiologically advantageous as more oxygen is delivered to the active regions.
(b)
Small mammals possess a much higher metabolic rate than humans and therefore it is appropriate that
oxygen should be released much more readily.
2
The position of the curve of the fetus relative to that of its mother means that its blood has a greater affinity for
oxygen than the maternal blood. This has to be so, as the fetus must obtain all of its oxygen from its mothers blood at the
placenta. So, at any given partial pressure of oxygen the fetal blood will take up oxygen from the maternal blood and will
always be more saturated with oxygen than the maternal blood. This is just as true for the human fetus.
3
This means that the blood has a high affinity for oxygen and that it is able to combine with it at the low oxygen
tensions experienced at high altitude. This is another good example physiological adaptation.
Control of ventilation (mechanism of breathing control)
Involuntary control of breathing is carried out by a breathing centre located in the medulla of the brain. The ventral
(lower) portion of the breathing centre acts to increase the rate and depth of inspiration and is called the inspiration centre.
The dorsal (top) and lateral (side) portions inhibit inspiration and stimulate expiration. These regions form the expiratory
centre. The breathing centre communicates with the intercostal muscles by way of the intercostal nerves and with the
diaphragm by way of the phrenic nerves. The bronchial tree (the mass of bronchioles and the bronchi) is connected to the
brain by the vagus nerve. Rhythmic nerve impulses to the diaphragm and intercostal muscles bring about ventilation
movements.
Inspiration inflates the lungs and, as they inflate, stretch receptors (also known as proprioceptors) located in the
bronchial tree are stimulated to send more and more nerve impulses via the vagus nerve to the expiratory centre. This
temporarily inhibits the inspiratory centre and inspiration. The external intercostal muscles therefore relax, elastic recoil of
the lung tissues occurs and expiration takes place. After this has occurred, the bronchial tree is no longer stretched and the
stretch receptors no longer stimulated. Therefore the expiratory centre becomes inactive and inspiration can begin again.
The whole cycle is repeated rhythmically throughout the life of the organism. Forcible expiration can be achieved by
contraction to the internal intercostal muscles.
A basic rhythm of breathing is maintained by the medulla even if all nervous input is cut. However, under normal
circumstances various stimuli modify this basic rhythm. The main stimulus that controls breathing is the concentration of
carbon dioxide in the blood, rather than the oxygen concentration. When carbon dioxide levels increase (as, for example,
during exercise) chemoreceptors in the carotid and aortic bodies of the blood system are stimulated and they send nerve
impulses to the inspiratory centre. The medulla itself also contains such chemoreceptors. The inspiratory centre then sends
out impulses via the eternal intercostal and phrenic nerves to external intercostal muscles and diaphragm causing them to
increase the rate at which they contract. This increases the rate of inspiration.
Control of ventilation in man
Ventilation of the respiratory system in man is primarily controlled by the breathing centre in a region of the hindbrain
called the medulla oblongata. The ventral portion of this centre controls inspiratory movements and is called the
inspiratory centre; the dorsal and lateral portions control breathing out and is called the expiratory centre. Control also
relies on chemoreceptors in the medulla, the carotid and aortic bodies of the blood system. These are sensitive to minute
changes in the concentration of carbon dioxide in the blood. When this level rises, increased ventilation of the respiratory
surfaces is required. Nerve impulses from these chemoreceptors stimulate the inspiratory centre in the medulla. Nerve
impulses from the inspiratory centre pass along the phrenic and external intercostal nerves to the diaphragm and external
intercostal muscles. Their increased rate of contraction causes faster inspiration. As the lungs expand, stretch receptors in
their walls are stimulated and impulses pass along the vagus nerve to the expiratory centre in the medulla. This
automatically switches off the inspiratory centre, the muscles relax and expiration takes place. The stretch receptors are
no longer stimulated, the expiratory centre is switched off and the inspiratory switched on. Inspiration takes place
again.
The breathing centre may also be stimulated by impulses from forebrain resulting in a conscious increase or
decrease in breathing.
The main stimulus for ventilation is therefore the change in carbon dioxide concentration and stimulation of
stretch receptors in the lungs; changes in oxygen concentration have relatively little effect.
Lung volumes and capacities
The average young female has a lung capacity of approximately 4 dm 3 and the average young adult male 5 dm3.

The tidal volume (VT) is the volume of air that enters and leaves the lungs at each natural resting breath (the
volume of gas exchange during one breath in and out).
The inspiratory reserve volume (IRV) is the volume of air that can be taken by a maximum inspiratory effort,
over and above the normal inspired tidal volume. In other words, this is the extra air that you can take in when
you breathe in as deep as possible after a normal inhalation
The expiratory reserve volume (ERV) is the volume of air that can be expelled by the most powerful expiratory
effort, over and above the normal expired tidal volume. This is the extra air breathed out when you force the air
out of your lungs as hard as possible after a normal expiration.
The vital capacity (VC) is the sum of the tidal volume and the inspiratory and expiratory reserves. It is the
volume of air which can be breathed out by the most vigorous possible expiratory effort following the deepest
possible inspiration.
The residual volume (RV) is the volume of air left in the lungs after the strongest possible exhalation. It has to be
measured indirectly.
The total lung capacity (TLC) is the sum of the vital capacity and the residual volume.

Stomata
Structure and functions
Stomata are pores in the epidermis through which gaseous exchange takes place. They are found mainly in leaves, but also
in stems. Each stoma is surrounded by two guard cells which surround a small pore a small pore a few microns wide
known as the stomatal aperture. Guard cells are different form normal epidermal cells in being kidney-shaped. Unlike
other epidermal cells, they possess chloroplasts and have denser cytoplasm with a more prominent nucleus. The guard
cells control the size of the stoma by changes in their turgidity.
The guard cell walls are unevenly thickened. the wall furthest from the pore (called the outer/dorsal) wall is
thinner than the wall next to the pore (the inner/ventral) wall. Also, the cellulose microfibrils that make up the walls are
arranged so that the inner wall is less elastic than the outer wall. Some of the cellulose microfibrils form hoops around the
kidney/sausage-shaped guard cells. These hoops are inelastic. As the cells inflate with water, that is become turgid, the
hoops tend to stop the cells increasing in diameter/width (getting fatter) so they can only expand by increasing in length.
Because the thin outer walls stretch more easily than the thick inner walls, each cell becomes semicircular in shape
(buckle outward). Thus a hole, the stoma, appears between the guard cells.
Mechanism of stomatal opening and closing
Starch-sugar hypothesis
During the day the chloroplasts of the guard cells produced sugars by photosynthesis. These made the water potential of the guard
cells more negative (lower) which therefore drew in water from adjacent cells by osmosis, causing increased pressure potential
and the opening of the stomata. When it was found that the opening occurred too rapidly to be accounted for in this way, a second
theory was substituted. It was thought that changes in the carbon dioxide concentration of leaves was responsible for stomatal
opening and closing. It could be observed that stomata opened when carbon dioxide concentrations were low and closed when it
was high. At high light intensities photosynthesis uses up carbon dioxide. Being acidic in solution, its removal causes a rise in pH
in the guard cells. This favours the conversion of starch to sugar by the enzyme starch phosphorylase. The sugar so produced
makes the water potential of the guard cells more negative (lower) which therefore take in water form the surrounding cells. The
resultant increase in pressure potential causes the stomata to open. The reverse situation occurs in the dark, when carbon dioxide
produced ins respiration causes a fall in pH. This theory cannot completely explain the process, however. In many plants the
changes in sugar concentration are inadequate to produce the necessary alteration in pressure potential. Furthermore, in some
species, like onion (Allium cepa), no starch at all is formed in guard cells.
K+ ions accumulation hypothesis
In the light (during the day), ATP is rapidly produced in guard cells, a potassium ion pump in the cell membranes actively
transports K+ ions from subsidiary cells (neighbouring epidermal cells) into the guard cells. K+ and malate ions accumulate in the
vacuole of guard cells, making the water potential of the guard cells more negative (lower). Water enters into guard cells by
osmosis, causing high hydrostatic pressure in these cells, i.e. guard cells become more turgid and effecting stomatal opening.
In the dark (at night), K+ ion pump in the cell membranes of the guard cells transfers K+ ions to the subsidiary cells. Ions are lost
from the guard cells; water moves out from the guard cells to the subsidiary cells, leading to lowered hydrostatic pressure in the
guard cells and effecting stomatal closing.

Stomata open and close in response to the movement of ions across the guard cells plasma membrane
The movement of H+ and K+ across the plasma membranes of guard cells explains the opening and closing of
stomata. Light, particularly blue light triggers the movement of potassium ions (K+) into the guard cells from
subsidiary cells or ordinary epidermal cells. This movement of K+ occurs by active transport through specific
5

ion channels in the guard cells plasma membranes and requires ATP. The ATP supplies energy to pump protons
(H+) out of the guard cells; the H+ is formed when malic acid produced in the chloroplasts of the guard cells
ionises. The removal of H+ from the guard cells produces an electrochemical gradient that can drive the uptake
of K+ through specific K+ channels. This ions movement is an example of a linked cotransport system. The K+
accumulates in the vacuoles of the guard cells.
The active uptake of K+ in the guard cells makes the water potential of guard cells to be more negative
(increase the solute concentration the vacuoles). As a result, water enters the guard cells from surrounding
epidermal cells by osmosis. The increased hydrostatic pressure (turgidity) of the guard cells changes their shape,
because the thickened inner cells walls do not expand as much as the outer walls. Thus, the stoma opens:
Light proton pump moves H+ out of guard cells K+ actively transported into guard cells guard
cells change shape and stoma opens
In the late afternoon or early evening, stomata close, possibly by a reversal of the opening process. The
K+ is pumped out of the guard cells into the surrounding epidermal calls. Water leaves the guard cells by
osmosis, the cells close their turgidity, and the pore closes as the guard cells collapse.