268 BIOLOGY
As you have read earlier, oxygen (O
2
) is utilised by the organisms to
indirectly break down simple molecules like glucose, amino acids, fatty
acids, etc., to derive energy to perform various activities. Carbon dioxide
(CO
2
) which is harmful is also released during the above catabolic
reactions. It is, therefore, evident that O
2
has to be continuously provided
to the cells and CO
2
produced by the cells have to be released out. This
process of exchange of O
2
from the atmosphere with CO
2
produced by the
cells is called breathing, commonly known as respiration. Place your
hands on your chest; you can feel the chest moving up and down. You
know that it is due to breathing. How do we breathe? The respiratory
organs and the mechanism of breathing are described in the following
sections of this chapter.
17.1 RESPIRATORY ORGANS
Mechanisms of breathing vary among different groups of animals
depending mainly on their habitats and levels of organisation. Lower
invertebrates like sponges, coelenterates, flatworms, etc., exchange O
2
with CO
2
by simple diffusion over their entire body surface. Earthworms
use their moist cuticle and insects have a network of tubes (tracheal
tubes) to transport atmospheric air within the body. Special vascularised
structures called gills (branchial respiration) are used by most of the
aquatic arthropods and molluscs whereas vascularised bags called lungs
(pulmonary respiration) are used by the terrestrial forms for the exchange
of gases. Among vertebrates, fishes use gills whereas amphibians, reptiles,
birds and mammals respire through lungs. Amphibians like frogs can
respire through their moist skin (cutaneous respiration) also.
B
REATHING AND
E
XCHANGE OF
G
ASES
C
HAPTER
17
17.1 Respiratory
Organs
17.2 Mechanism of
Breathing
17.3 Exchange of
Gases
17.4 Transport of
Gases
17.5 Regulation of
Respiration
17.6 Disorders of
Respiratory
System
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BREATHING AND EXCHANGE OF GASES
17.1.1 Human Respiratory System
We have a pair of external nostrils opening out above the upper lips.
It leads to a nasal chamber through the nasal passage. The nasal
chamber opens into the pharynx, a portion of which is the common
passage for food and air. The pharynx opens through the larynx region
into the trachea. Larynx is a cartilaginous box which helps in sound
production and hence called the sound box. During swallowing glottis
can be covered by a thin elastic cartilaginous flap called epiglottis to
prevent the entry of food into the larynx. Trachea is a straight tube
extending up to the mid-thoracic cavity, which divides at the level of
5th thoracic vertebra into a right and left primary bronchi. Each bronchi
undergoes repeated divisions to form the secondary and tertiary bronchi
and bronchioles ending up in very thin terminal bronchioles. The
tracheae, primary, secondary and tertiary bronchi, and initial
bronchioles are supported by incomplete cartilaginous rings. Each
terminal bronchiole gives rise to a number of very thin, irregular-walled
and vascularised bag-like structures called alveoli. The branching
network of bronchi, bronchioles and alveoli comprise the lungs (Figure
17.1). We have two lungs which are covered by a double layered pleura,
with pleural fluid between them. It reduces friction on the lung-surface.
The outer pleural membrane is in close contact with the thoracic
Bronchus
Lung
heart
Diaphragm
Epiglottis
Larynx
Trachea
Cut end of rib
Pleural membranes
Alveoli
Pleural fluid
Bronchiole
Figure 17.1 Diagrammatic view of human respiratory system (sectional view of
the left lung is also shown)
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270 BIOLOGY
lining whereas the inner pleural membrane is in contact with the lung
surface. The part starting with the external nostrils up to the terminal
bronchioles constitute the conducting part whereas the alveoli and their
ducts form the respiratory or exchange part of the respiratory system.
The conducting part transports the atmospheric air to the alveoli, clears
it from foreign particles, humidifies and also brings the air to body
temperature. Exchange part is the site of actual diffusion of O
2
and CO
2
between blood and atmospheric air.
The lungs are situated in the thoracic chamber which is anatomically
an air-tight chamber. The thoracic chamber is formed dorsally by the
vertebral column, ventrally by the sternum, laterally by the ribs and on
the lower side by the dome-shaped diaphragm. The anatomical setup of
lungs in thorax is such that any change in the volume of the thoracic
cavity will be reflected in the lung (pulmonary) cavity. Such an
arrangement is essential for breathing, as we cannot directly alter the
pulmonary volume.
Respiration involves the following steps:
(i) Breathing or pulmonary ventilation by which atmospheric air
is drawn in and CO
2
rich alveolar air is released out.
(ii) Diffusion of gases (O
2
and CO
2
) across alveolar membrane.
(iii) Transport of gases by the blood.
(iv) Diffusion of O
2
and CO
2
between blood and tissues.
(v) Utilisation of O
2
by the cells for catabolic reactions and resultant
release of CO
2
(cellular respiration as dealt in the Chapter 14).
17.2 MECHANISM OF BREATHING
Breathing involves two stages : inspiration during which atmospheric
air is drawn in and expiration by which the alveolar air is released out.
The movement of air into and out of the lungs is carried out by creating a
pressure gradient between the lungs and the atmosphere. Inspiration
can occur if the pressure within the lungs (intra-pulmonary pressure) is
less than the atmospheric pressure, i.e., there is a negative pressure in
the lungs with respect to atmospheric pressure. Similarly, expiration takes
place when the intra-pulmonary pressure is higher than the atmospheric
pressure. The diaphragm and a specialised set of muscles – external and
internal intercostals between the ribs, help in generation of such gradients.
Inspiration is initiated by the contraction of diaphragm which increases
the volume of thoracic chamber in the antero-posterior axis. The
contraction of external inter-costal muscles lifts up the ribs and the
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BREATHING AND EXCHANGE OF GASES
sternum causing an increase in the volume of
the thoracic chamber in the dorso-ventral axis.
The overall increase in the thoracic volume
causes a similar increase in pulmonary
volume. An increase in pulmonary volume
decreases the intra-pulmonary pressure to less
than the atmospheric pressure which forces
the air from outside to move into the lungs,
i.e., inspiration (Figure 17.2a). Relaxation of
the diaphragm and the inter-costal muscles
returns the diaphragm and sternum to their
normal positions and reduce the thoracic
volume and thereby the pulmonary volume.
This leads to an increase in intra-pulmonary
pressure to slightly above the atmospheric
pressure causing the expulsion of air from the
lungs, i.e., expiration (Figure 17.2b). We have
the ability to increase the strength of
inspiration and expiration with the help of
additional muscles in the abdomen. On an
average, a healthy human breathes 12-16
times/minute. The volume of air involved in
breathing movements can be estimated by
using a spirometer which helps in clinical
assessment of pulmonary functions.
17.2.1 Respiratory Volumes and
Capacities
Tidal Volume (TV): Volume of air inspired or
expired during a normal respiration. It is
approx. 500 mL., i.e., a healthy man can
inspire or expire approximately 6000 to 8000
mL of air per minute.
Inspiratory Reserve Volume (IRV):
Additional volume of air, a person can inspire
by a forcible inspiration. This averages 2500
mL to 3000 mL.
Expiratory Reserve Volume (ERV):
Additional volume of air, a person can expire
by a forcible expiration. This averages 1000
mL to 1100 mL.
Figure 17.2 Mechanism of breathing showing :
(a) inspiration (b) expiration
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272 BIOLOGY
Residual Volume (RV): Volume of air remaining in the lungs even after a
forcible expiration. This averages 1100 mL to 1200 mL.
By adding up a few respiratory volumes described above, one can
derive various pulmonary capacities, which can be used in clinical
diagnosis.
Inspiratory Capacity (IC): Total volume of air a person can inspire
after a normal expiration. This includes tidal volume and inspiratory
reserve volume ( TV+IRV).
Expiratory Capacity (EC): Total volume of air a person can expire after
a normal inspiration. This includes tidal volume and expiratory reserve
volume (TV+ERV).
Functional Residual Capacity (FRC): Volume of air that will remain in
the lungs after a normal expiration. This includes ERV+RV.
Vital Capacity (VC): The maximum volume of air a person can breathe in
after a forced expiration. This includes ERV, TV and IRV or the maximum
volume of air a person can breathe out after a forced inspiration.
Total Lung Capacity (TLC): Total volume of air accommodated in the
lungs at the end of a forced inspiration. This includes RV, ERV, TV and
IRV or vital capacity + residual volume.
17.3 EXCHANGE OF GASES
Alveoli are the primary sites of exchange of gases. Exchange of gases also
occur between blood and tissues. O
2
and CO
2
are exchanged in these
sites by simple diffusion mainly based on pressure/concentration
gradient. Solubility of the gases as well as the thickness of the membranes
involved in diffusion are also some important factors that can affect the
rate of diffusion.
Pressure contributed by an individual gas in a mixture of gases is
called partial pressure and is represented as pO
2
for oxygen and pCO
2
for
carbon dioxide. Partial pressures of these two gases in the atmospheric
air and the two sites of diffusion are given in Table 17.1 and in
Figure 17.3. The data given in the table clearly indicates a concentration
gradient for oxygen from alveoli to blood and blood to tissues. Similarly,
TABLE 17.1 Partial Pressures (in mm Hg) of Oxygen and Carbon dioxide at Different
Parts Involved in Diffusion in Comparison to those in Atmosphere
Respiratory Atmospheric Alveoli Blood Blood Tissues
Gas
Air (Deoxygenated) (Oxygenated)
O
2
159 104 40 95 40
CO
2
0.3 40 45 40 45
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BREATHING AND EXCHANGE OF GASES
a gradient is present for CO
2
in the opposite direction, i.e., from tissues to
blood and blood to alveoli. As the solubility of CO
2
is 20-25 times higher
than that of O
2
, the amount of CO
2
that can diffuse through the diffusion
membrane per unit difference in partial pressure is much higher compared
to that of O
2
. The diffusion membrane
is made up of three major layers
(Figure 17.4) namely, the thin squamous
epithelium of alveoli, the endothelium of
alveolar capillaries and the basement
substance (composed of a thin basement
membrane supporting the squamous
epithelium and the basement membrane
surrounding the single layer endothelial
cells of capillaries) in between them.
However, its total thickness is much less
than a millimetre. Therefore, all the factors
in our body are favourable for diffusion of
O
2
from alveoli to tissues and that of CO
2
from tissues to alveoli.
Figure 17.4 A Diagram of a section of an
alveolus with a pulmonary
capillary.
Figure 17.3 Diagrammatic representation of exchange of gases at the alveolus and
the body tissues with blood and transport of oxygen and carbon dioxide
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274 BIOLOGY
17.4 TRANSPORT OF GASES
Blood is the medium of transport for O
2
and CO
2
. About 97 per cent of O
2
is
transported by RBCs in the blood. The remaining 3 per cent of O
2
is carried
in a dissolved state through the plasma. Nearly 20-25 per cent of CO
2
is
transported by RBCs whereas 70 per cent of it is carried as bicarbonate.
About 7 per cent of CO
2
is carried in a dissolved state through plasma.
17.4.1 Transport of Oxygen
Haemoglobin is a red coloured iron containing pigment present in the
RBCs. O
2
can bind with haemoglobin in a reversible manner to form
oxyhaemoglobin. Each haemoglobin molecule can carry a maximum of
four molecules of O
2
. Binding of oxygen with haemoglobin is primarily
related to partial pressure of O
2
. Partial pressure of CO
2
, hydrogen ion
concentration and temperature are the other factors which can interfere
with this binding. A sigmoid curve is obtained when percentage saturation
of haemoglobin with O
2
is plotted against the
pO
2
. This curve is called the Oxygen
dissociation curve (Figure 17.5) and is highly
useful in studying the effect of factors like
pCO
2
, H
+
concentration, etc., on binding of O
2
with haemoglobin. In the alveoli, where there
is high pO
2
, low pCO
2
, lesser H
+
concentration
and lower temperature, the factors are
all favourable for the formation of
oxyhaemoglobin, whereas in the tissues,
where low pO
2
, high pCO
2
, high H
+
concentration and higher temperature exist,
the conditions are favourable for dissociation
of oxygen from the oxyhaemoglobin. This
clearly indicates that O
2
gets bound to
haemoglobin in the lung surface and gets
dissociated at the tissues. Every 100 ml of
oxygenated blood can deliver around 5 ml of
O
2
to the tissues under normal physiological
conditions.
17.4.2 Transport of Carbon dioxide
CO
2
is carried by haemoglobin as carbamino-haemoglobin (about
20-25 per cent). This binding is related to the partial pressure of CO
2
.
pO
2
is a major factor which could affect this binding. When pCO
2
is high
and pO
2
is low as in the tissues, more binding of carbon dioxide occurs
whereas, when the pCO
2
is low and pO
2
is high as in the alveoli, dissociation
20
0
20
40
40
60
60
80
80
100
100
Partial pressure of oxygen (mm Hg)
Percentage saturation of haemoglobin with oxygen
Figure 17.5 Oxygen dissociation curve
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BREATHING AND EXCHANGE OF GASES
of CO
2
from carbamino-haemoglobin takes place, i.e., CO
2
which is bound
to haemoglobin from the tissues is delivered at the alveoli. RBCs contain
a very high concentration of the enzyme, carbonic anhydrase and minute
quantities of the same is present in the plasma too. This enzyme facilitates
the following reaction in both directions.
CO H O H CO
Carbonic
anhydrase
Carbonic
anhydra
2 2 2 3
+
→
←
sse
HCO H
→
←
+
− +
3
At the tissue site where partial pressure of CO
2
is high due to
catabolism, CO
2
diffuses into blood (RBCs and plasma) and forms HCO
3
–
and H
+,
. At the alveolar site where pCO
2
is low, the reaction proceeds in
the opposite direction leading to the formation of CO
2
and H
2
O. Thus,
CO
2
trapped as bicarbonate at the tissue level and transported to the
alveoli is released out as CO
2
(Figure 17.4). Every 100 ml of deoxygenated
blood delivers approximately 4 ml of CO
2
to the alveoli.
17.5 REGULATION OF RESPIRATION
Human beings have a significant ability to maintain and moderate the
respiratory rhythm to suit the demands of the body tissues. This is done
by the neural system. A specialised centre present in the medulla region
of the brain called respiratory rhythm centre is primarily responsible for
this regulation. Another centre present in the pons region of the brain
called pneumotaxic centre can moderate the functions of the respiratory
rhythm centre. Neural signal from this centre can reduce the duration of
inspiration and thereby alter the respiratory rate. A chemosensitive area
is situated adjacent to the rhythm centre which is highly sensitive to CO
2
and hydrogen ions. Increase in these substances can activate this centre,
which in turn can signal the rhythm centre to make necessary adjustments
in the respiratory process by which these substances can be eliminated.
Receptors associated with aortic arch and carotid artery also can recognise
changes in CO
2
and H
+
concentration and send necessary signals to the
rhythm centre for remedial actions. The role of oxygen in the regulation of
respiratory rhythm is quite insignificant.
17.6 DISORDERS OF RESPIRATORY
SYSTEM
Asthma is a difficulty in breathing causing wheezing due to inflammation
of bronchi and bronchioles.
Emphysema is a chronic disorder in which alveolar walls are damaged
due to which respiratory surface is decreased. One of the major causes of
this is cigarette smoking.
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276 BIOLOGY
SUMMARY
Cells utilise oxygen for metabolism and produce energy along with substances
like carbon dioxide which is harmful. Animals have evolved different mechanisms
for the transport of oxygen to the cells and for the removal of carbon dioxide from
there. We have a well developed respiratory system comprising two lungs and
associated air passages to perform this function.
The first step in respiration is breathing by which atmospheric air is taken in
(inspiration) and the alveolar air is released out (expiration). Exchange of O
2
and
CO
2
between deoxygenated blood and alveoli, transport of these gases throughout
the body by blood, exchange of O
2
and CO
2
between the oxygenated blood and
tissues and utilisation of O
2
by the cells (cellular respiration) are the other steps
involved.
Inspiration and expiration are carried out by creating pressure gradients
between the atmosphere and the alveoli with the help of specialised muscles –
intercostals and diaphragm. Volumes of air involved in these activities can be
estimated with the help of spirometer and are of clinical significance.
Exchange of O
2
and CO
2
at the alveoli and tissues occur by diffusion. Rate of
diffusion is dependent on the partial pressure gradients of O
2
(pO
2
) and CO
2
(pCO
2
),
their solubility as well as the thickness of the diffusion surface. These factors in
our body facilitate diffusion of O
2
from the alveoli to the deoxygenated blood as
well as from the oxygenated blood to the tissues. The factors are favourable for the
diffusion of CO
2
in the opposite direction, i.e., from tissues to alveoli.
Oxygen is transported mainly as oxyhaemoglobin. In the alveoli where pO
2
is
higher, O
2
gets bound to haemoglobin which is easily dissociated at the tissues
where pO
2
is low and pCO
2
and H
+
concentration are high. Nearly 70 per cent of
carbon dioxide is transported as bicarbonate (HCO
3
–
) with the help of the enzyme
carbonic anhydrase. 20-25 per cent of carbon dioxide is carried by haemoglobin
as carbamino-haemoglobin. In the tissues where pCO
2
is high, it gets bound to
blood whereas in the alveoli where pCO
2
is low and pO
2
is high, it gets removed
from the blood.
Respiratory rhythm is maintained by the respiratory centre in the medulla
region of brain. A pneumotaxic centre in the pons region of the brain and a
chemosensitive area in the medulla can alter respiratory mechanism.
Occupational Respiratory Disorders: In certain industries, especially
those involving grinding or stone-breaking, so much dust is produced
that the defense mechanism of the body cannot fully cope with the
situation. Long exposure can give rise to inflammation leading to fibrosis
(proliferation of fibrous tissues) and thus causing serious lung damage.
Workers in such industries should wear protective masks.
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BREATHING AND EXCHANGE OF GASES
EXERCISES
1. Define vital capacity. What is its significance?
2. State the volume of air remaining in the lungs after a normal breathing.
3. Diffusion of gases occurs in the alveolar region only and not in the other parts of
respiratory system. Why?
4. What are the major transport mechanisms for CO
2
? Explain.
5. What will be the pO
2
and pCO
2
in the atmospheric air compared to those in the
alveolar air ?
(i) pO
2
lesser, pCO
2
higher
(ii) pO
2
higher, pCO
2
lesser
(iii) pO
2
higher, pCO
2
higher
(iv) pO
2
lesser, pCO
2
lesser
6. Explain the process of inspiration under normal conditions.
7. How is respiration regulated?
8. What is the effect of pCO
2
on oxygen transport?
9. What happens to the respiratory process in a man going up a hill?
10. What is the site of gaseous exchange in an insect?
11. Define oxygen dissociation curve. Can you suggest any reason for its sigmoidal
pattern?
12. Have you heard about hypoxia? Try to gather information about it, and discuss
with your friends.
13. Distinguish between
(a) IRV and ERV
(b) Inspiratory capacity and Expiratory capacity.
(c) Vital capacity and Total lung capacity.
14. What is Tidal volume? Find out the Tidal volume (approximate value) for a healthy
human in an hour.
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