UNIT RESPIRATION

Introduction
Objectives
Respiratory Gases
Partial Ressure
Sqlubility .
Diffusion of Gases
Modes of Respiration
Gills
Ventilation of Gills
Gas Exchange
Lungs
Mammalian Lungs
Regulation of Respiration
Adaptations for Diving and Underwater Swimmivg
Tracheae ,
Transport of Gases in ~ l o o d
Haemoglobin
Oxygen Transport in Blood
Carbon Dioxide Transport in Blood
Summary
Terminal Questions
Answers
< -
2.1 INTRODUCTION
You have learnt in Unit 1 that virtually all animals depend on oxidation of.food
materials for their energy requirements. They utilise oxygen and produce carbon
dioxide during the course of their metabolism. The process of oxygen uptake and
release of carbon dioxide is called respiration and applies to hole organisms as well
d"
-as to the processes that take place in the cells: You have come aquainted with the
'dynamics of cellular respirario-n in LSE-01. This unit focuses on the acquisition of
oxygen 'by animals from the environment' and its distributioh to cells where it is
needed for cellular respiration. You will also learn how the same distribution system
'eliminates carbbn dioxide or the metatlolic waste. Animals takeup oxygen from the .
medium they live ih. Aquatic animals take up oxigen from water; terfestrial animals
from air.
'

To start with, we have explained"some 6f the phys'lcal properties of gases. It is

important that you compreheid the mainepoints because they ate essential for your
understanding of respiratory physiology. To enter or leave the body, oxygen and
carbon dioxide must cross a barrier of living tissue: In simplest animals gas exchange
, takes pla& through the skin or through plasma membrane (as-in unicellular
organism), but most multic+lular animals have more elaborate respiratory
mechanisms which ensure that blood and respiratory medium are brought in close.
contact over a large respiratory surface. ow ever, the most'suc+ssful respirat~ry
strategy is one that suitsthe medium in which thd animal lives. We will deal with
three contrasting mechariisms for respiration. Teleost gills, as eqamples of organs that
take up sxygen dissolved in water, mammalian lungs as examples of air breathing
organs and insect tracheal system as example of uptake and cirwl.ation-of gaiseS'
. withaul the involvement of a circulatory system. The movement of respiratory
medium into and out of the respiratory organswill need to be regulated particularly
when we note that the demand for gxygen, increases occasionally (as in exercise).
We will discuss this control brielly. '
. .
. . . --

.Later
. in the iinit you will also l e m - a b o u t r&@ratory pigments.that aid in the
transport.of oxygen. . . .. . . . . . . - . ~.
A . J., :--* '
You would notice tbat in our discussion,& the respiratory process we will, refer to
the circulatory system quite often for it is +te difficult to discuss one sydem with
the exclusion of all others. As you go tkrough'the course you will realise that in
animals all systerqs' function in an integratedma2ner and no system functions alone. .
 In the next unit we will discuss circulatory syst& in detail where we will weave in
findings from this unit again.

Objectives
After a careful study of this unit you will be able to :
explain the need for gas exchange between the environment and respiratory
surfaces in animals
describe the structural and functional difference between gills, lungs and trachae,
giving their advantages and disadvantages
'
explain how countercurrent exchange mechanism works in the gills of bony fish
and state the importance of this adaptation.
I
explain the process of gas transport between the blood and tissues in mammals.
discuss the role of haemoglobin in transfer of oxygen and compare it to the
transport of carbon dioxide in blood.

2.2 RESPIRATORY GASES

Physiologically the most important gases are oxygen, carbon dioxide and nitrogen.
Pa (Pascal) is the SI unit for Molecular oxygen comprises 21% of the atmosphere, carbon dioxide is only 0.3% .
pressure. In this unit we have still
used the rnrn v g to denote
while nitrogen forms 78.0%. A11 the gases present in the air exert a combined
pressure as a I'arge body of pressure which is referred to as one atmosphere (760 mm Hg) or 101.3 k Pa.
knowledge stili exists in the older
unit. And rnrn Hg is still used to Most of the oxygen is in the air but some of it is also dissolved in the bodies of water
denote pressure in respiratory and and in soil water. An animal's immediate source of oxygen or respiratory medium
circulatory functions. , therefore, is either water or air. Comparison of the gaseous composition and physical
characteristics of air and water emphasise the adaptations needed by terrestrial and
aquatic animals to overcome the problems of respiration. For example, the oxygen
content of air is about 20 times that of water saturated with air. The diffusion rate
for oxygen in air is much more than the diffusion rate in water. In addition there is
another hazard. Carbon dioxide diffuses rapidly from air into water. Therefore,
elimination of carbon dioxide in water and air are different. ,
x -.
Extraction of oxygen from different media thus presents special problems as a result
of the physical characteristics of the environment. Therefore, to understand the
physiology of respiration, we must have a basic knowledge of some of the physical
properties of gases. The exchange of oxygen and carbon dioxide either in the
respiratory organ or at the tissue level is dependent on the partial pressure,
concentration and diffusion of gases. Let us consider them one by one.

2.2.1- Partial Pressure

The behaviour of an individual gas in a mixture (for example, the rate at which
oxygen diffuses across a respiratory membrane exposed to air) depends largely on
/
.
the partial pressure p of that gas.
Imagine a gas filled chamber containing different gases, each gas exerts the pressure
that it would exert if it was alone in that space. Therefore, the total pressure P in
that chamber will be a sum of the pressures exerted by all the gases in the mixture.
The partial pressure of each gas in the mixture is proportional to the volume of that
gas within the total volume of the gas mixture. So, we can calculate partial pressure as:

where p is the partial pressure of the gas

x is the percentage of the specific gas in the total volume
P is the total pressure of the mixture (101.3 k Pa or 760 mm H$, for dry air).
Thus partial pressure for oxygen at sea level would be

and for carbon dioxide the value would be

When we calculate the partial pressure of a gas in air we must remember that
atmospheric air is not always dry. It usually contains water vapour which contributes
to the total pressure. The amount of water vapour varies with temperature. For
instance at 37OC and 760 mm Hg the partial pressure of water vapour - P w r is about
47.26'mm Hg. Under these conditions gases contribute only 712.74 mm Hg of the
total pressure. Therefore, equation (1) must be modified to accommodate water
vapour.

In respiratory functions equation (2) is more applicable because at the interphase of

the respiratory lining water vapour will show a gradient directed towards the airand
water would be lost in that direction.

2.2.2 Solubility
Most of us know by experience
You are ;ware that gases are soluble in liquid. Solubility of a gas in a liquid depends that solubility decreases with
on the partial pressure, temperature and presence of other solutes. Table 2.1 shows increasing temperature. When
the solubility of oxygen, carbon dioxide and nitrogen at different temperatures. You you put water for boiling, the
can see that solubility of carbon dioxide is much greater than oxygen, and that as the bubbles formed at the sides
temperature increases, solubility decreases. before boiling are gases escaping
out of the solution as the
temperature rises.
Table 2.1 :Solubility of gases in cm30f ges per c m h f water

Temperature - Gas '

)I
("C) c o 2 0 2 N2

It is also important to know that solubility of gases decreases with the presence ot
solutes or increased salinity. Table 2.2 shows the solubility of oxygen at various
temperatures in freshwater and seawater.

Table 2.2 : Effect of temperature on the amount of oxygen dissolved in fresh water
and seawater in equilibrium with atmospheric air

Temperature Freshwater (ml 0, Seawater (mi 0,

C'c) per litre of water) per litre of water)

2.2.3 Diffusion of Gases

Gases diffuse from areas of higher partial pressure to areas of lower partial pressure.
The diffusion along the partial pressure gradient ceases only when the partial pressure
in both areas becomes equal. This is true in aquous solutions: within mixtures of gases
and across gas-water interphase. Very seldom, diffusion in direction of partial
-
'pressure gradient also means diffusion along a conce;ltration gradient.
_-_- - --- Ld

In later sections we shall see how the movement of oxygenand carbon dioxide
depends on the paltial pressure gradient \hat exists at theinterphase between vascular
:fluibs and air in lubgs. Within the body &atthe capillarjt level the intracellular fluids
and vascular fluids develop opposed partial pressure gradients and hence gases move
- -

opposite directions.
 Having studied the physical basis of gas diffusion we can now proceed t o see how
animals exchange gases across their respiratory surfaces. First, we shall discuss
various arrangements for respiratory gas exchange in animals.

SAQ 1
a) Pick out the correct statements from the following:
i) Partial pressure of a gas in solution is defined as the partial pressure of that
gas in atmosphere in equilibrium with the solution.
ii) As temperature decreases solubility of gases decreases too.
iii) As the partial pressure of a gas inwa water sample is reduced, the gas tends
to move out of the solution.
iv) The diffusion of a gas along a partial pressure gradient is always along the
concentration gradient.
b) Suppose air at 30°C and 735.18 mm Hg is saturated with water vapour at 18.00
mm Hg. What would the partial pressure of oxygen be in this saturated air?

2.3 MODES OF RESPIRATION

Many small organisms obtain oxygen by diffusion through their body surfaces. They
do not have any specialised respiratory organs nor do they have blood circulation.
Larger and more complex organisms, however, need specialised surfaces for gas
exchange and a circulatory system that transports oxygen more readily than that
possible by simple diffusion.

Calculations based on metabolic demands and rate of diffusion in protoplasm %how

Jelly fish can be very large. Since that simple diffusion is sufficient only to meet the demands of organisms not larger*
it has less than 1% organic matter
and the red 99%water and salt, it than 1 mm in diameter. These calculations appear reasonable when we see that
has a very low average oxygen animals like protozoans and flatworms that meet their respiratory requirements
consumption rate. The actively through diffusion are either quite small or have very low metabolic rates. Giant land
rnetabolising cells are located planarians may be 50 cm long but they are flat with very large surfaces in relation t o
near the surface therefore
diffusion alone is sufficient for gas
mass, therefore, diffusion is sufficient to meet their oxygen demand. Coelentrates,
exchange. corals and sponges often reach very large sizes but have very modest metabolic
demands. Sponges and corals maintain a circulation of water by cilia over the surfaces
of cells which line their canal systems. Thus sufficient gas exchange takes place
without the aid of a circulatory system or respiratory pigments

In animals that have efficient circulatory systems and readily permeable vascular
skins, gas exchange occurs through the integument. Thus we find that animals like
earthworms, leeches and many larval fishes are among the many animals that obtain
the oxygen they need across their general body surface. Even larger animals such as
many amphibians and fish may rely on cutaneous respiration during emergencies or
use it as a supplement to the gills or lungs.

The ability to respire through the integument developed the most in eels, and
amphibians that have moist and highly vascularised skins. The integumentary
contribution to respiration may be as low as 20 per cent in dry-skinned toads to 76
per cent in the urodele Triturus alpestris and over 90 per cent in the giant salamander
Cryptobranchus alleganiensis. This aquatic amphibian is the largest animal relying
exclusively on integumentary breathing. The adults are 25 to 60 cm in length and may
weigh over 1 kg. These animals.1ack gills and the lungs being unspecialised play little
or no part in respiration.
Animals that are large and have higher metabolic rates have specialised respiratory
organs. These organs have a thin respiratory surface to help in gas exchange. (Fig. 2.1)
Respiratory
,
organs may be of the following types:
Those that have respiratory surface turned out forming an evagination. These are
called gills.
Those that have respiratory surface turned in forming an invagination. These are
called lungs. Our own lungs are good examples.

(d) Trachc

Fig, 2.1 :Some aquatic animals and animals that maintain moist skins, can excbnge gaws directly through
their integuments (a), which are in contact with water. Gills (b) highly folded external tissues that
-
provide enormous surfaces for exchanging gases with water are common among larger aquatic
animals. Terrestrial animals that exchange gases with the atmosphere do so by means of highly
branched internal surfaces that are protected from drying out. Many vertebrates have sac-like
lungs (c): insects and some other arthropods have systems of branched tubes called tracheae (d).

Insects have a special respiratory system. Small openings on the insect's body surface
connected to small tubes or trachea that branch and spread throughout the body. The
gases diffuse through these branches directly into the cells.
Generally gills are for aquatic breathing and lungs for breathing air. We shall now
consider specific kinds of respiratory organs employed by animals that live in water
and on land.
SAQ 2
Why does diffusion alone sufficktd supply oxygen in both Paramecium and jelly fish?
:,.. 1
................................................................................................................

2.4 GILLS
Gills are highly vascularised extensions of gas exchange membranes. The s i m ~*<$:
type may be extention fo ce as in s e a . s l u P s . r R and manv ot'
 slow moving aquatic animals. 'The complexity of the gills reflect the animal's
requitieaht for oxygen. Animals with greater need for oxygen have elaborate gill
stcucture. Before we discuss the structure of the gill let us consider the ways in which
gills are ventilated because the environment in contact with respiratory surface must
be continuously renewed to avoid depletion of oxygen or excessive &cumulation of
carbon d i o x s

2.4.1 Ventilation of G%
Various mechanical devices are usedio'increase the flow of water over the gill's
surface. There can be two means of increasing ihe flo-water over the gill surface:
--
i) by movement of gills as seen in sm>il o r p s m s . Some aquatic insect larvae use
this method, but this is not a very efficient and practical method. The force
needed to overcome resistance to movement is great and the energy needed to
move the gill also increases correspondingly. The mechanical strength of the gill
.. . in most larger animals would also need to be increased.

ii) by moving water over the gill. This is achieved by ciliary action as in gills of
mussels and clams. Movement of water due to a mechanical pump is a device
used by fishes and crabs. The locomotion of many aquatic animals .helps to
circulate the water. Many pelagic fish, especially tuna, swim rapidly through the
water and create a rapid flow over the gills. Among invertebrates the octopus or
squid ventilates its gills by taking water in the mantle cavity and ejecting it out
through the siphon. This jet propulsion also provides the locomotion.

2.4.2 Gas Exchange

Gill surface area must be large enough to provide adequate exchange of gases.
Therefore, highly active fish have largest relative gill area. Fig. 2.2 compares highly
active fish with sluggish bottom dwellers. Their relative surface area is also given.

Mackerel - 2551 Eel - 902

puffer -505 Goosefish - 51

Fig. 2.2 :Fast sw~mmingfish like makerel have larger gill area than sluggish.bottom dwelling fish. The total
surface area of gills is expressed per unit per gram body weight of fish.

For gas exchange to be adequate there should be adequate flow and close contact
between the gill and water. T o understand how this happens let us look at the
structure of a gill in bony fishes as a representative of an aquatic respiratory surface.
Gills are enclosed in a gill cavity. This provides protection for the fragile organ and
also permits the water to run over the gills in an efficient manner. Gills of fishes
' consist of several gill arthes on either side. The gill arches separate the o ercular and
the buccal cavities. From each gill afch extend two rows of gill filaments. Fig. 2.3 P
shows the arrangement of gill filaments in the arches. The tips of the filaments of
adjacent arches meet forming Z-sieve like structure through which the water flows.
 rl.J~'
-
BUCCAL , Muscle
CAVITY /

Fig. 2.3 : a) Position of gill arches beneath the operculum on thehft side of fish. The operculurn has been
lifted to show the arch.
b) Part of two'adjoining gill arches with their fdaments. Note that the tips meet to form a sieve
like arrangement for flow of water. The water moves through the mouth over the branched
gills. Solid arrows show the flow of water.
C) Part of a single Nament showing the flat e e l i a e ; the flow of water is opposite to the direction
in which the blood moves.

Each filament has an upper and a lower row of flat lamellae. The lamellae of
successive filaments are in close contact. Gas exchange takes place in these lamellae
a s water flikvs between them in one direction and blood within them in the other
direction. Thispi5Tall a countercurrent flow. This type of flow has an important
consequence. I$ permits the fish gills to have the highest possible oxygen levels.
+/

'&5-10
2 1 2 4 - 9
- 6
- 3
- 3
t o
2 \

9
(a) wuntercurrent

4
I /
&s-12 - 9 4 6 3

t . P O - 2 4 -6-
r 0 t l
. (b) concurrent
t -
Fig. 2.4 :~ouhtercurrentt~0~7ijfavou6bettiro~~~en
- -
a=rption by blood. The numbers in the figure
C A
-indicatetht?p@M pressure of 0, (Po3 in water and blood. Blood enters with low Po2but leaves
the l a d h e with nearly the same POz as water; while water looses most of its O2before leaving.
Countercurrent flow also reduces the energy eqst'at pumping watel; over the gills.
b) If blood and water moved in the same direction i.e. conwrrent flow, then no further e x c h g e
of Oz would take place after equal concentration is reached. Then the Po2 in Mood would at
themost reach the Poz levels of the outflowing water.

Fig. 2.4 shows the advantage of countercurrent flow. You must remember (a) that
diffusiotl of oxy5en from water to blood depknds largely on the steepness of the
- -- - -
 gradient of partial pressure of oxygen between the two compartments; (b) the most
efficient gas exchange system is one that ensures the highest possible partial pressure
of oxygen in the blood leaving the gill. Now let us suppose that the incoming blood
is devoid of all oxygen, and imagine the flow of water and blood is in the same
direction i.e. it is concurrent (Fig. 2.4 b) when the two streams come in contact first
there is a steep pressure gradient from the wker t o blood. Oxygen is transferred from
the water to blood at a high rate till an equilibrium is reached after which no transfer
occurs. Now consider Fig. 2.4a which shows countercurrent flow. When blood which
has zero Po2 comes in contact with water for the first time, the water also has low
Po2 (since it has been loosing oxygen on its way to this point) but still sufficient for
a pressure gradient to be maintained. As the blood moves on it meets with water
richer in oxygen and the Po2 of blood increases steadily. At all points along the
capillary, Po2 gradient is sufficiently high to permit transfer of oxygen from water to
blood. The net effect is that blood leaving the gills in countercurrent exchange has
extracted 80 per cent or more of the dissolved oxygen from water.
To move water over the gills, teleosts use combined pumping.action of mouth and
opercular cover. Water is drawn into the mouth, passes over the gills and flows out
through the opercular clefts, valves guard the entrance to the buccal cavity and
opercular clefts ensuring a unidirectional flow of water (Fig. 2.5). The volume of the
buccal cavity can be changed by lowering of the jaw and the floor of the mouth and
that of the opercular cavity is changed by the movements of opercular flaps that swing
out to enlarge the cavity and swing in to reduce it. Changes in volume of both cavities
is synchronised but a pressure differential is maintained across the gills throughout
the breathing cycle.
The pressure in the opercular cavity is always slightly lower than the pressure in the
buccal cavity providing for a continuous flow of water.
Buccal Opercular
Cavity Cavity
Gills I
1 I

I I + I

< Operculum
1
.<- Closed

Increasing Decreas~ng
Volume vd;rne
Fig. 2.5 : Dual pump in fish provides a continuous unidirectional flow of water.
3
Some fish do not use pumping action for gill ventilation. It has been known for long
that large tunas cannot be kept alive in captivity unless they are put in circular tanks
where they can swim continuously. The fish swim with their mouths partly open and
there are no visible breathing movements. Water flows continuously over their gills.
This is known as ram ventilation.
It is now known that many fish breathe by pumping at low speed and change to ranl
ventilation at high speed. In this case the work of breathing is transferred from the
muscles of the opercular pump to the muscles of the body and tail. However, ram
ventilation is more economical as far as energy consumption is concerned than
opercular pumps at high rates required for fast swimming.
SAQ 3
Most fish when taken out of water become asphyxiated, although there is far more
oxygen in the air thanqin water. Apart from the reason that the weight of the gills
cannot be supported by them in air, what other factor is responsible for this.
You have seen that respiration in water can be explained through some simple
physical principles. Let us now move on to examine respiratory gas exchange in
animals that breathe air.

2.5 LUNGS
In section 2.2, you studied that air has more oxygen than.water. The atmosphere
provides a consiant supply of oxygen almost everywhere. The greatest disadvantage
however, is that air tends to dry out gas exchange membranes. To overcome this,
terrestrial animals have to live in extremely moist environments.or find some ways
to keep their respiratory surfaces moist. The water loss is minimised by turning the
respiratory membranes inside the body into lungs in amphibians, reptiles, birds and
mammals, and trachea in insects.

Before we proceed to describe terrestrial respiration by lungs let us examine some

major advantages of breathing air. If we compare water and air we find that water
in equilibrium with atmosphere at 15°C contains only 7 ml of oxygen per litre. In
contrast 1 litre of air contains 209 ml of oxygen. Water is much more viscous than
air, therefore, aquatic organisms have to expend wore energy to move the medium
over the respiratory surface. Apart from this oxygen diffuses some 10,000 times more
rapidly in air than in water and so can diffuse in lungs over several millimeters while
diffusion from water to respiratory surface in fish gill can take place in only minute
fraction of a millimeter. The flow of oxygen into the respiratory cavity in terrestrial
animals is well regulated according to the oxygen demands of the organism.

Lungs can be simple, characterised by air exchange with surrounding environment by

diffusion only. These are called the diffusion lungs and are present in small animals
such as pulmonate snails, small scorpions, s a n e spiders and some isopods. The other
type -ventilation lungs are typical of vertebrates. The air passes through a tube into
inflatable lungs where gas exchange takes place and oxygen poor, carbon dioxide rich
air is then forced out usually through the same tube. This is known as tidal flow of
air. Ventilation of the lungs can take place in two different ways:

1) By using a pressure pump as in amphibians. Fig. 2.6 shows the process of

ventilation in frog. The inflation of lungs depends on positive pressure
bucco-pharengeal pump. The nares remain open while glottis is closed (the air
does not enters the lungs). The &or of the buccal cavity is raised and lowered
periodically (Fig. 2.6a). At irregular intervals the glottis is open and nares are
closed. The floor of the buccal cavity is raised forcing air into the lungs. As a
result the frog can take in air several times without exhaling and blow itself up
to considerable size. The glottis can close and while the air remains inside the
lungs the cycle is replated in the buccal cavity.

2) By using a suction pump. Exhalation can be passive and inhalation is aided by

musele contraction or as in mammals, by contraction of muscular dome shaped
diaphragm and external intercostal muscles lifting the ribcage. This decreases the
pressure in the pleural space s o causing the lungs to expand and air flows in.

2.5.1 Mammalian Lungs Fig. 2.6 : Breathing cycle in the

frog
In this unit we will study mainly mammalian lung as it is the best representative of a a) dris'takenintothebueepl
respiratory surface adapted for terrestrial respiration. For this purpose, human lung cavity by lowering it.
can be taken as a model as shown in Fig. 2.7. b) drispermlttcdtoeacapeltan
the lmgs passing over tbe
buccal cavity.
When we breathe, the air enters the wind pipe or trachea which are divided into right c) tbeeXternalll~resarecIQ8Cd
and left bronchi (Fig. 2.7). These in turn branch repeatedly 'forming bronchioles. The air is forced into the lungs.
fine branches of the bronchioles lead into alveolar sacs, which are clusters of minute d) whik alr 1pnuio8in lungs. Tbe
cycle an be rejIe8te.d.
sacs, whose diameter ranges between 150 to 300 micron or micrometer. f i e alveoli
have thin walls and capillaries from the pulmonary artery extensiyely occupy the
vascular side of the alveoli (Fig. 2.7). A pair of human lungs contain about 300 million
alveoli and the total surface area is about 70 meter square. This area is nearly equal
 Blood
-
Bronchioles
. noy ,

Pleural '
space

Fig.9.7 :In liumans and other mammals, plr pssss~eaIndude nostrils, nasal cavity (and mouth), pharynx,
larynx, bechea, bmebl, and bmehides. The bmchidea termhate In numerous Mind alveoll
(see inset). The spongy lungs are subdivided into lobes, with tbree in the right long and two in the
/
left lung. The muscular, shelf-llke diaphragm seals the thoracic cavity di from the abdominal
cavify below.

Measurement of surface of frog to a tennis court! Gas exchange takes.place only in the alveoli. Trachea, bronchi and
lung indicates that 1 cm3 lung theirbranches are only connecting tubes. When we breathe out, these tubes are filled
tissue has total g a exchange
sunface of 20 cm2 per 1 cm3 of
with used air from the lungs and when we breathe in again this used air is pushed
lung tissue only. ss shows that back into the lungs first, before the fresh air enters. The volume of air in the passage
large surface area is related to high thus reduces the volume of fresh air that can enter the lungs. Therefore, this space
metabolic rate of wann blooded is called anatomical dead space. The dead space volume is 150 cm3. The volume of
mimais. air inhaled in one breath is called the tidal volume and for a normal man at rest it is 1
about 500 cm3. Therefore, only 350 cm3 (500 - 150 = 350) of fresh air reaches the
alveoli. In exercise the dead space volume is not significant. For example, a man may
breathe 3000 cm3 of air in a single breath, in that case 150 cm3 is hardly significant. 1
1
No matter how hard one may try, the lungs are never empty of air, There is still
1650 cm3of air left in it. This is known as residual volume. During inhalation, 350 (50@
150) cm3 of fresh air enters and this mixes with the 1650 cm3 of air already in the
lungs. The renewal of air is only one part in five. Therefore, the lungs have a constant
composition of 15% oxygen and 5% carbon dioxide. This remains same even during
exercise as kcreased ventilation during exercise supplies the extra oxygen
requirements of the body.

In Fig. 2.8 you can see the thin lining of alveolar capillary membrane which is 0.2 to
0.6 thick. Gas exchange actually takes place here. Wherever there is exchange of
gases, the barrier must be thin and it is so in the alveolar membrane. This membrane
separates vascular fluidsfand alveolar space. The inner space of the alveoli which
comes in contact with the inspired air is covered with alveolar epithelium. This
epithelial surface is lined by a lipoprotein mixture called lung surfactant. This surface
lining reduces the surface tension at interphase, across which opposed diffusion
gfadients of oxygen and carbon dioxide are developed. Another function of the
surfactant is to help in the first breath taken by an infant or by a new born animal.
Before birth the alveoli are collapsed and the thin layer of moisture between them
makes it difficult for them-to be opened (try opening a plastic bag with a thin film of
water between it). Surfactant reduces the muscular effort to open the lungs.
 Capillary Alveolar Respiration
endothelial iceH epithelial cell

Capillary
endothelium
(0.04-0.2 pm)
\, )-- Erythrocyte
(0.75~)

Alveolo-

(0.2-0.6crm)
I
surface ~ntektitial Alveolar
lining space epithelium
(0.01r m ) (0.02-0.2 p m ) (0.05-0.3 pm)
I
1 ALVEOLAR SPACE (50-300,pm) 1
Fig. 2.8 :Microscopic structure of respiratory lining of human lung.
a) Note the presence of smooth muscles that can haven marked effect on the dimension of the air
ways.
b) Higher rnagnifkatlon of alveolmpillary membrane which separatk alveolar space from
capillary blood.
8

Exchange in Alveoli
The blood that enters the lungs from the heart has been routed through body tissue
where mitochondria1 respiration, (refer to LSE-01, Units 11 and 12) has depleted its
oxygen content. It thus has a low partial pressure of oxygen (Po2 = about 40 mm
Hg). At the same time metabolic activity increases partial pressure of carbon dioxide
in body tissue and partial pressure of carbon dioxide of blodd entering the alveoli is
45 mm Hg.
So compared to partial pressures in atmosphere blood entering lungs from the body
tissue has low Po2 and high Pco2. The partial pressure of oxygen in alveoli is high
and that of carbon dioxide is low so the gases diffuse only along their partial pressure
gradient. The whole process of exchange is depicted in Fig. 2.9.
Atmospheric air

7%. 2.9 :Exchaoge of respiratory gases in lungs and tissue ceb. Numbers represent yartial pressures in
millimeters of mercury (mm Hg).
 A~I~UIP~--I 2.5.2 Regulation of Respiration
Whenever the need for oxygen in the body increases, ventilation of the respiratory
organs also increases. In the same way whenever the level of oxygen in the medium
falls, the ventilation must increase or the body should be able to extract more oxygen
from the respired airpor both these processes must take place.
In lungs of warm-blooded animals i.e. birds and mammals the ventilation is regulated
primarily by the amount of carbon dioxide in the lung air. If we add more carbon
dioxide to the inhaled air, there is a rapid increase in ventilation, even if carbon
I
dioxide content in inhaled air is increased to that found normally in the lungs (5%),
.- respiratory ventilation volume increases several folds. In higher concentrations,
carbon dioxide becomes dangerous. Oxygen on the other hand, has a much smaller
tov effect on ventilation, if we reduce the oxygen concentration from 21% to 18.5% there
is virtually po effect.
The rythmic contractions of the diaphragm and of the interrmstal muscles are
controlled by a respiratory centre which is located in the area of the medulla -
oblongata and pons of the brain. There are separate neurons for inspiration and
I
expiration that work alternatively.
The respiratory c q t r e j s sensitive to increased carbon dioxide levels and increased
acidity of the blood. It is peculiar that even though metabolism requires oxygen yet
the respiratory centre is not very sensitive to decreased oxygen levels. However,
chemo-receptors located in carotid bodies and aortic arches near the heart respond
Heart)
to decreased Po2 levels by increasing the depth and frequency of breathing. Fig. 2.10
Fig. 2.10: AetIm of carbon dioxide
depicts the control process graphically.
receptor miis In contrast aquatic animals that breathe through gills are very sensitive to oxygen
levels. ~espiratorymovement in many specie? increases if oxygen levels in
surrounding me$ium fall. In amphibia the respiration in tadpoles is regulated by
' t
oxygen levels and in adult
* frogs the mechanism is controlled by carbon dioxide levels.
SAQ4 ,
3
Mark True or False among the following statements:
i) Lungsurf+ctankincreases the s u r f ~ etension of the alveoli so that they don't
collapsl.
ii) Tidal voluhz is the volume of air taken in one breath.
iii) .Anatomical dead spac. 1s the volume of afr present in the trachea, broncheoles
etc.
-%
iv) ~esidual-airin the rungs is important for maintaining the carbon dioxideJevels
in lung.
~ l ~controlled by q part of the brain called cerebrum.
v) Breathrr:, , . ~ , ~ o r : l r ; iis
+'.
2.5.3 Adaptations for Diving and Underwater Swimming .
ManITfTZfs'ana birds cannot breathe when swimming underwater, but ca'rbon dioxide
continues to be brought to the lungs. As carbon dioxide builds up in the alveolar air,
the usuat Sthnulus for breathing is given and the underwater swimmer has to come
to thesurmce to breathe. Many species of mgmmals for example otter, beavers, seal'
and whak show adaptations which permit them to stay underwater. a

s seen best in the sperm whale that dives as deep as lOOd meters
There a d a p r g i ~ nare
for over an hour without any respiratory distress. Physiology of diving is an
interesting subject but it is beyond the scope of this unit to discuss it in detail. Briefly
we can mention that diving mammals have to conserve their oxygen for as long as
possible. For this they have evolved certain adaptations. They are :
i) The heart rate is slowed down along with a fall in blood supply to muscles, skin.
and visera so that the flow to the brain and heart is maintained. In some mammals
receptors in the nasal region when stimulated by water, trigger cardiac slowing..
ii) Diving vertebrate, also have relatively large blood volumes and venous reservoirs.
iii) During a deep dive the high pressure of the water causes the lungs to collapse
partially and thus capillary circulation is also restricted there by reducing the
solutjon of air in the blood.
 Unlike other mammals that have successfully adapted to underwater, human
beings are limited by their respiratory physiology. Thus we see that human divers
face several physiological problems. The major one is the increased ambient
pressure underwater. For every 10 meters there is an increase by 1 atmosphere
(101.3 k Pa) i.e. if one dives 10 m below sea level the pressure is doubled and
the amounts of gases dissolved in plasma would also double. The increased levels
of oxygen and nitrogen in blood can have serious consequences. Oxygen toxicity
develops rapidly when thesPo2 rises above 2.5 atmosphere because of oxidation
of enzymes and other destructive changes that can damage the nervous system
and lead to death. For this reason deep sea divers use a mixture of gases instead
of pure oxygen. Large amounts of nitrogen dissolved in blood under pressure
causes nitrogen narcosis, a condition resembling*intoxication. Another danger
arises when divers surface too rapidly. The gases expand and their bubbles form
emboli in the circulatory system.
Human beings swimming underwater at shallow depths can endanger themselves if
they hyperventilate or overbreathe. They can ihus reduce the carbon dioxide levels
from the alveoli. This will delay the onset of carbon dioxide-respiratory drive and the
sensation of the need to breathe. But meanwhile oxygen will continue to be used up
and since Po2 receptors are not as sensitive as Pcoz receptors the divers may become
unconscious through lack of oxygen and thus drown.

2.6 TRACHEAE
Insects that have successfully colonised the terrestrial'environment have envolved a
respiratory system very different from other land animals - the tracheal system.
I Apart from insects other arthropods, such as millipedes, centipedes, a few archinids
l and terresrrial crustaceans havc tracheae as respiratory organs.
The tracheal system consists of an air filled system of tubes -the tracheal tubes that
branch repeatedly until they become fine as capillaries, and are called tracheoles. The
tracheolar endings arc in close proximity to the cells of the body and sometimes they
indent into the plasma membrane giving the impression that they actually enter the
cells. Tracheac vary in diameter from 1-2 mm and the diameter of tracheoles is
between 0.6-0.8 mm. In the larger tracheae, thickcnings known as taenidia in the
cuticular lining prevent the tubes from collapsing. The walls of the tracheae become
progressively thinner as thcy branch so that the tracheolar endings are only 5,urn in
dianictcr. Fig. 2.1 1 shows the nrrangc~ncntof trachci~lsystcm in some insccts.

Air s;~cs

Pig. 2.1 1 :Some modifications of the tracheal system in insects. (a) The basic pattern, (b) mechanically
ventilated air sacs, (c) terminal spiracles in aquatic insects.

The tracheae open to the outside by structures known as spiracles. Usually there are
10 pairs of spiricles. They have closing mechanisms, aided by valves, muscles and
filters. Such closing mechanisms also help to conserve water. The tracheoles and
tracheolar endings are permeable to water and may contain variable amounts of fluid. 43
Animal Physiology -I The fluid enters by capillary action from the cells. At rest the tracheolar endings are
filled with fluid but during activity the cell contents become hypertonic due to
accumulation of metabolites and the fluid is withdrawn into the cells from the
tracheolar endings. This increases the much needed oxygen supply to the actively
metabolising tissue.

Ventilation of Tracheal System

The manner in which tracheae are ventilated varies with species of insects but
movement of air is achieved in two ways :
1) Diffusion
2) Passive Suction Ventilation
1) Diffusion : Gaseous exchange due to diffusion generally takes care of the oxygen
needs of most small insects. Similarly fairly large but inactive insects (e.g., some
large caterpillars) do not actively ventilate the system. Diffusion alone is
adequate provided the spiracles are kept open permanently. The same process
accounts for elimination of carbon dioxide.
Passive Suction Ventilation : Many active insects and insects that live in
environments where water is scarce cannot depena on diffusion alone to obtain
their oxygen. Flying insects need a steady supply of oxygen, therefore, their flight
muscles are richly supplied with tracheoles. If the spiracles of such insects were
to be kept open they would loose all their body water. To solve this problem
there is control of volume of air and the direction of flow through the system.
For example locusts maintain a flow of air through the spiracles by muscular
contraction and accurately timed opening and closing of various spiracles. The
air enters through the anterior spiracles and leaves by the posterior spiracles
establishing a unidirection flow of air. However, gas exchange in the tracheoles
is through diffusion only even in-the largest and most active insects.
Another .very interesting system of spiracular control is seen in some insect species.
The spiracles are kept closed tightly most of the time. When the insect requires
oxygen the spiracles open partially and close rapidly again. This 'fluttering' of the
spiracle lets the air enter in a streaming movement without much loss of water. Once
a high partial pressure of oxygen is reached the spiracles close.
While the oxygen is utilised by the tissues the carbon dioxide is stored temporarily
in the tissues as bicarbonates thus keeping the Pcoz within tolerable limits, when Pco,
reaches a critical limit the spiracles open fully and expel the carbon dioxide in a single
burst and the cycle is repeated.
The efficiency of ventilation is greatly improved in certain species by presence of air
sacs. These are balloon like dilation of the trachae (Fig. 2.11 b). Their formation and
deflation depends on haemolymph pressure and muscular contractions of the body.
Fig. 2 . 1 1 ~shows a modification found in some aquatic insects. Most of the spiracle$
are non-functional. 'Jnly the two last ones open to the outside. They are so located
that the insect cdn make periodic excursions to the surface and make contact with
the atmosphere for oxygen either by diffusion or by respiratory movements. The
mosquito larvae is a common example.
In many species of aquatic insects the air in the trachae is replenished by diffusion of
oxygen via tracheal gills. Many aquatic insects carry a temporary store of air as a
bubble on the body surface which replenishes the air in tracheae during submergence.
In other aquatic insects a constant volume of air is held as a thin layer close to the
body surface by water repellent hair forming a plasteron (or layer of air). As long as
water is well aerated insects with plasteron can remain submerged for many months.
As much as one-third or half of the total capacity of the air system can be emptied
in one expiration giving a renewal of about half the volume of the respiratory system.
This is much greater than in a mammal at rest, where in each breath fifth
of the air contained in the system is renewed. ->

SAQ 5
a) If a stream of carbon dioxide is directed towards the spiracles of an insect, what
would be the most perceptible effect?
 b) How does pouring kerosene in open drains or stagnant water pools prevents
a breeding of mosquitoes?

2.7 TRANSPORT OF GASES IN BLOOD

We know that two types of gas exchanges are constantly occurring in the animal
body--one at the interphase of the respiratory membrane and external environment
and the other in the rest of the animal tissues. The principle underlying is the same
at both places - passive diffusion along a pressure gradient. I

In many invertebrates oxygen and carbon dioxide are carried dissolved in blood or
haemolymph. The amount of oxygen carried in simple solution is small, therefore, in
highly organised animals (many invertebrates-and all vertebrates) oxygen is
transported bound reversibly to proteins that are oxygen carriers. These proteins
contain a metal, commonly iron or copper and are coloured. They are known as
respiratory pigments. The common respiratory pigments are listed in Table 2.3 along
with their molecular weights and occurrence in the animal kingdom.
Table 2.3 : common respiratory pigments, their properties and occurrence in the animal kingdom

Pigment Colour Site Molecular Oxygen Animal

Weight Volume%
Haemocynin Blue Plasma 3-6.8 X lo6 1-5 Molluscs
(copper containing) 4-8 x lo5 1-5 Crustaceans
Haemerythrin Red Corpuscles 6.6 x lo4 2 Annelids
(iron containing)
Chlorocruorin Green Plasma 3.4 x lo6 9 Annelids
(iron containing)
Haemoglobin Red Corpuscles 6.8 x lo4 15-30 Mammals
(iron containing) 10-25 Birds
7-12 Reptiles
6-13 Amphibians
1-16 Gnathostome
fishes
1.7 x lo4 1.2 Cyclostome
fishes
Plasma 3.0 x lo6 5-15 Annelids
1.5 x loh 1-6 Molluscs

2.7.1 Haemoglobin
Among the respiratory pigments we shall consider haemoglobin in some detail as this
is the most familiar, widespread as well as the most efficient respiratory pigment. You
can see from Table 2.3 that more efficient haemoglobins are those that are packed
in cells and combine with far greater amounts of oxygen. Fig. 2.12

PO, =I00 mm Hg O2 1 Oxyhaemoglobin

Fig. 2.12 :Plasma and whole blood U~atare brought into equilibrium with the same gas mixture have Ute
same Po, and therefore, the same amount of dissolved oxygen molecules (shown as black dots).
The oxygen content of whole blood however, is much more because haemoglobln binds to oxyged
-rlnn..ln.
 If the amount of haemoglobin that is packed in cells was to be free in plasma, the
viscosity of blood would be like syrup. Apart from this, being enclosed in cells
provides a more stable environment as reaction of oxygen and haemoglobin is
affected by ion concentration and organic compounds in blood. .
The structure of haemoglobin was briefly mentioned in Unit 5 of the course in Cell
Biology (LSE-01). Let us now consider the structure in more detail.
A group of compounds called porphyrins are widely distributed in plants, animals
and bacteria. Porphyrins associate with metals to form metalloporphyrins which form
a variety of compounds. For example chlorophyll about which you will read in
Block-3 is a magnesium-porphyrin complex and cytochromes which contain
porphyrin group play an important role in intracellular oxidation.
When ferrous iron Fef + is added to a porphyrin protoporphyrin IX, ferrous
porphyrin or haem is formed. The molecular structure is shown in Fig. 2.13.

CH, CH=CH,

HC-C
I I
C=CH
II "N' I I
-C I
I
C =C - CH,
\
N -----Fe -N '
I
1

CH2
I
COOH

I
COOH

Fig. 2.13 :Chemical structure of haem group (a), Mematic representative of a single submit of haemoglobii(b).

The ferrous iron atom is bound to the four nitrogens of the protoporphyrin ring. The
ferrous iron makes two more links, one with the oxygen atom to form.
oxyhaemoglobin and the other to the globin portion of the molecule at a single amino
acid sit& (Fig. 2.13b).
The haem group of all haemoglobins is the same and the variations in amino acid
sequences of the globin group form the different haemoglobins.
Myoglobin may also have an We can build up the structure of haemoglobin by considering a single unit or
oxygen storage function especially monomeric form caned myoglobin. This consists of a single polypeptide chain the
.useful in the functioning of heart. globin in which the haem group is embedded. Myoglobin is found in striated muscles
During diastole when coronary
blood flow is maximum it stores of vertebrates andcombines with one molecule of oxygen. The vertebrate
oxygen and when during systole haemoglobins are tetramers formed by the aggregation of four polypeptide chains
the coronary arteries are each containing a discrete haem group. Thus each haemoglobin can combine with
squeezed the stored oxygen is four molecules of oxygen forming oxyhaeomoglobin in a reversible reaction and the
released.
unoxygenated compound is called deoxyhaemoglobin. Two monomers in human
haemoglobin are of a type called s and the other two are P type. You would recall
the unique quaternary structure of haemoglobin from Unit-5 LSE-01.
Nearly all vertebrate haemoglobins are tetramers but invertebrate haemoglobins are
more diverse. The most distinctive feature is that the subunits often form large
aggregates of relative high molecular weights (Refer t o Table 2.3).
In mammalian blood the amount of physically dissolved oxygen is about 0.2 ml of
oxygen per 100 ml of blood. The amount found bouild to haemoglobin is 20 ml
oxyeen per 100 ml of blood. The dissolved oxygen is therefore, almost insignificant
in animals without respiratory pigments. The only exception is the antarctic fish which
 lacks a respiratory pigment altogether. The explanation could be that at low
temperature, the metabolic rate is low and oxygen like other gases has higher
solubility at low temperature.

1 2.7.2 Oxygen Transport in Blood

I
All four respiratory pigments are adapted to load and unload oxygen effectively in
the habitats where they have evolved, whether animals live on land where the air
contains 210 ml of oxygen per litre or in fresh water containing 8.0 mlnitre or in the
sea containing 6.4 mltlitre. The loading unloading reaction can be written.as
respiratory surface
(haemoglobin) Hb + OZ%============A
tissue
Hb 02(oxyhaemoglobin)

The direction of this reaction depends on Po2 of the environment and bond strength Haemoglobin has even a stronger
or affinity between haemoglobin and oxygen. In the lungs where Po, is high almost affinity for carbon monoxide than
oxygen. The bond is 210 times
all deoxyhaemoglobin molecules bind to oxygen. Low Po2 in the systemic capillaries stronger. Carbon monoxide tends
promotes unloading. Similarly strong affinity favours loading and weak bonding

I
to displace oxygen in
favours unloading. Haemoglobin has a bond strength which permits 97% of haemoglobin and remains
haemoglobin to combine with oxygen when leaving the lung. At the same time the attached as the blood passes
bond is sufficiently weak to permit unloading in tissues. Under normal resting through the tissues. ~ h transport
k
of oxygen is impaired leading to
conditions 22% of the oxygen is unloaded. This satisfies the oxygen need of the body dangerous consequences and even
simultaneously maintaining a reserve that is utilised during emergency conditions. death.

Oxygen Dissociation Curves

The oxygen content of blood fully saturated or oxygenated can b;e calculated. It is
known as the oxygen capacity of blood and this varies for different species. In humans
the oxygen carrying capacity is 20 ml oxygen per 100 ml blood. The relationship of
oxygen carrying capacity to surrounding oxygen concentration can be shown
graphically by oxygen dissociation curves. These curves are obtained by subjecting
blood samples to different partial pressures of oxygen. The per cent oxyhaemoglobin
saturation at different partial pressures of oxygen are plotted.
The oxygen dissociation curve (shown in Fig. 2.14) is S-Shaped or a sigmoid curve.
From the graph we can see how haemoglobin acts as a carrier of oxygen. Total
saturation occurs in the lungs where arterial pressure is above 95 mm Hg and the
oxygen is unloaded at low Po2 found in tissues (about'40 mm Hg).

20
15
I
Amount of 0, unloaded
to tissues

Fig. 2.14 :Oxygen dissociation curve shaws how haemoglobin's oxygen binding capacity depends on partiel
. pressure of oxygen. Note that there is a 22% decrease in per cent oxyhaemoglobin as blood passes
from arteries to veins in tissue. This results in ranioading of approximately 5 ml of oxygen per
100 ml of b i d .

Also the curve shows that changes in Po2 values from arterial to venous blood result
in 97-75 = 22% unloading when resting. During exercise this unloading is increased
to 39%. Fig. 2.15 shows the dissociation curve for myoglobin yhich in contrast is
rectangular. Myoglobin is described as middleman in the transfer of,oxygen from
blood to mitochondria within muscle cells. This is known as facilitated diffusion of
oxygen.
 Animal Physiology -I
I
I
I

I
I
I
I
I
I
I
I
I
I
I
I
I

Venous blood Arterial blood

I
I
I

Partial pressure of oxygen (millimeters of mercury)

Fig. 2.15 :A comparison of the dissociation curves for haemoglobin and for myoglobin. At the Poz of venous
blood, the myoglobin retains almost all of its oxygen, indicating a higher aff~nitythan haemoglobin
for oxygen. The myoglobin does, however, release its oxygen at the very low Poz values found
inside the mitochondria.
'
Fig. 2.15 also indicates that the compound remains oxygenated until quite low levels
of Po2 in the surrounding fluids is reached. You can see from the figure that at
20 mm Hg the haemoglobin in the blood is about 30% saturated but the myoglobin
in the muscles is above 80% saturated.
In molecular terms how does a S-shaped curve arise? Look at the part of the curve
in Fig. 2.14 where Po, is low. At first the slope is not steep but rises steeply up to a
certain point (about 20 mm Hg).
In other words haemoglobin does not take up oxygen at low Po, but as the
oxygenation of the pigment occurs its affinity for more oxygen increases. No such
increase in affinity is evident from the myoglbbin curve shown in Fig. 2.15. In
myoglobin the haem reacts to oxygen independently. In the case of haemoglobin
where 4 subunits are present, acquisition of one molecule of oxygen increases the
affinity of neighbouring haems for oxygen. This 'is known as co-operativity between
active sites. Let us see how this cooperation between active sites modifies the binding
of oxygen. It is now known that the haemoglobin can occur in two.interchangeable
forms, one is called T (tense) structure and the other is R (released) structure.
T structure has relatively low affinity for oxygen and R structure has high affinity for
oxygen. The uptake of oxygen by one subunit in T structure changes the whole
complex to R structure, Which then binds to oxygen one hundred times faster than
the first haerh group.
Oxygen dissociation curve for a sample of blood is affected by several factors. The
most important of them are :
In fever or exercise there is a 1) Temperature
higher rate of oxygen
consumption. Therefore, it is 2) pH
advantageous that haemoglobin 3) co2
delivers 0, more readily at high
4) Organic Phosphates
I temperatures.
At higher temperature haemoglobin gives up oxygen more readily and the
dissociation curve shifts to the right. This is of physiological importance because
increased temperature means higher metabolic rate or higher oxygen requirement.
Another important influence is the pH. Increase in carbon dioxide or other acids
lowers the pH of plasma and shifts the dissociation curve to the right (Fi,g. 2.16). A t
high carbon dioxide concentrations more oxygen is given up at any given oxygen
pressure. This effect is known as Bohr effect after the Danish scientist who first
described it. Therefore, as carbon dioxide enters the blood from respiring tissues it
encourages the release of more oxygen. This is an important characteristic because
it allows more oxygen to be released to tissue which need it the most. More oxygen
is loaded in the lungs and more unloaded at the tissues due to Bohr effect than would
be the case if only diffusion along the concentr'ation gradient was responsible.
 1 1 I I I

2 0 4 0 6 0 8 0 1 0 0
Partial pressure of 0, in mm Hg

(a)

Fig. 2.16 :Bohr effect. Oxyhaemoglobin surrenders its oxygen more readily in the presence of increasing
acidity. (a) Tbe oxygen dissodatlon curve shifts to the right with increasing acldity;
(b) rtabolkally active cells will receive more oxygen than metabolkally less active cells.

Carbon dioxide lowers the oxygen affinity of haemoglobin even if the pH-is kept
constant. This effect is due to the binding of carbon dioxide to the term~nalamino
groups of haemoglobin molecule. This site is not the same site on the molecule where
oxygen is bound.
The presence of organic phosphates in the red blood cells helps to explain many
peculiarities of the oxygen dissociation curve. Previously the red blood corpuscle was
~ ~ i ~ ~ * ~ ~ " ' - ~ ; ~ s p ~ n o
considered to be a bag full of haemoglobin with no metabolism of its own because of in blood bank,ng. Old
the absence of a nucleus. Now we know that it has.an active carbohydrate metabolism stored red cells loose then ability
and RBC has high content of ATP and 2,3-diphosphoglycerate (DPG). 2,3-DPG is a to produce 2.3-DPG which means
product of glycolysis and it binds to P chain of the globin and reduces oxygen affinity. that such cells will not unload
their oxygen easily. Modem
Experimentally it was shown that pure haemoglobin has greater oxygen affinity than techniques for storage of blood,
whole blood (the dissociation curve for pure haemoglobin is far to the left of the therefore,include the addition of
curve for whole blood). If 2,3-DPG is added to pure haemoglobin solution the oxygen energy substrates for resplrat~on
affinity decreases and approaches that of whole blood. The effect of 2,3-DPG is also and phosphate sources needed for
important in transfer of oxygen from maternal to foetal blood. Foetal haemoglobin of 213-DPG
is different from adult haemoglobin in having 2 6 chains instead of P chains. Foetal
haemoglobin therefore cannot bind to 2,3-DPG and thus has higher affinity for
oxygen at a given Po2. This higher oxygm affinity in foetal blood facilitates c -oen
transfer from mother to foetus.
SAQ 6
Pick out the true statements from those given below :
i) The haemoglobin of adults contain four identical subunits and a haem grc
ii) When haem and globin fractions are separated the haem group binds, reversibly
with oxygen.
iii) Bohr's effect facilitates transfer of oxygen to the tissues because of increased C 0 2
levels in blood.
iv) 2,3-DPG inhibits the transfer of O2 across the foetal-maternal barrier.
v) In haemoglobin one subunit after attaching to an oxygen facilitates the oxygen
binding to others.

2.7.3 Carbon Dioxide Transport in Blood

The same transport system that brings oxygen to the tissues must take back carbon
dioxide to the environment across the respiratory surface. However, unlike oxygen
that is transparted exclusively by haemoglobin, carbon dioxide is transported in three
ways:
1) as dissolved carbon dioxide in plasma (about 8%) and in red cell.
2) as carbaminohaemoglobin. About 25% of the total blood carbon dioxide is
camed attached to the amino groups in haemoglobin.

Haemoglobin -N '4H 4 + C 0 2 + $I++

Haemoglobin - NHCOO-
.
 It is then carried to the lungs where haemoglobin releases it in exchangebfor
oxygen.
3) as carbonic acid and bicarbonate which accounts for most of the carbon dioxide
carried by blood.
Carbon dioxide combines with watel; to form carbonic acid.
C 0 2 + H 2 0 + H2CO3
This reaction occurs spontaneously in plasma at a very slow rate but much more
rapidly within the blood cell due to the catalytic reaction of an enzyme, carbonic
snhydrase. The formation of carbonic acid is favoured by high Pco2 in the capillaries
of tissues. Carbonic acid dissociates rapidly in the red blood cells into hydrogen ion
(H+) and bicarbonate iog (HCO 7).
H2C03 + H + + HCO?
The H+ released are buffered by their combination with haemoglobin and HC0;
moves out of the cells. The inside of the cell thus gains a net positive charge. This
attracts chloride ion (CI-) which move inside the red blood cells. This exchange of
anions as the blood moves through capillaries in tissue is known as chloride shift (Fig.
2.17a). The red blood cells are very permeable to both Cl- and HCO; because the
membrane has a high concentration of a special anion carrier protein called band 111
protein that binds CI- and HCO; and transfers them-in opposite direction through
the membrane.

Plasma Red blood cells Capillary

(a)
I

Fig. 2.17 :Carbon dioxide trpnsport in blood (a)"in c M d e shift, serbon dioxide is transported in three
e,
forms: as d b l v e d CO, attached to haemoglobha s ~ i n o h a e m o g l o b i nand
, M crrbbnic
acid and blearbonate, (b) carbon dloxklc is r e l d - f r o m the Mood rr it Levels t h
capillaries. A reverse chloride shift occurs and carbonic acid is transformed into CO, and HzO.
 The formation of carbonic acid enhances oxygen unloading (Bohr effect) and oxygen
unloading in turn improves the ability of blood t o form carbonic acid and transport
carbon dioxide. When blood reaches pulmonary capillaries deoxyhaemoglobin is
converted to oxyhaemoglobin which has a lower affinity for H+. The H+ is released
in the red blood cells. This attracts HCO; from plasma which combines with H + to
form H2C03
H+ + HCO, --+ H2C03
Under low Pco2 of pulmonary vessels carbonic anhydrase catalyses the formation of
carbonic acid to carbon dioxide and water
Carbonic anhydrase
H2C03 Low Pco,
H2O + C02
Therefore, a reverse chloride shift occurs in the pulmonary capillaries to convert'
carbonic acid, and biocarbonate to C 0 2 gas which is eliminated in expired breath
(Fig. 2.17b).

After having discussed the properties of blood and its role in the transport of ,
respiratory gases we can now proceed to discuss the movement of blood in animals
in the next unit on circulation.

2.8 SUMMARY

.
In this unit you have studied that :
The physiological basis of gas exchange in animals depends on some simple
physical principles like solubility of gases, their diffusion rates which in turn
depends on the nature of gas, its partial pressure, temperature and presence of
other solutes.
The four basic categories of animal gas exchange mechanisms are diffusion through
body surface, gills, lungs and trachea. The design of respiratory surface and the
mechanism of breathing are related to the nature of the medium in which the
animals live.
Most aquatic animals exchange gases through gills that are evaginations of
respiratory surfaces. Fish gills consist of gill arches with rows of gill filaments. They
contain extensive capillary beds in plate-like lamellae. Water flow across the
lamellae opposes blood flow, setting up an efficient countercurrent exchange.
Diffusion of oxygen in water is slow, therefore, aquatic animals must expend
energy to move large volumes of water over the respiratory surfaces.
The vertebrate lung is a highly branched inpocketing of respiratory surface
containing numerous air sacs or alveoli, intimately associated with capillaries and
ventilated by a tidal flow of air.
Gas exchange in lungs takes place in alveoli where partial pressure of oxygen is
higher than that in blood hence oxygen diffuses from alveoli to blood. In body
tissues the partial pressure of oxygen is low hence oxvgen diffuses from blood into
tissues. Respiration is regulated by levels of Pcoz and by a respiratory centre in
the brain which is sensitive to blood Pco2.
Insects have evolved a separate system for gas exchange-tracheal system
independent of circulatory system.
Respiratory gases are transported mainly by respiratory pigments in blood. The
most well-known pigment is haemoglobin which combines with oxygen to form
oxyhaemoglobin. Oxygen dissociation curve depicts per cent oxyhaemoglobin
saturation at different values'of Po2. Carbon dioxide, pH, temperature and
presence of organic compounds in blood influence this cun;e. A fall in pH
decreases the oxygen affinity of haemoglobin for oxygen. This is Bohr effect.
Decreased affinity causes unloading of oxygen to tissue. Oxygen affinity also
decreases in the presence of 2,3-DPG.
Carbon dioxide is transported to the lungs by formation of carbonic acid in red
blobd cells. This reaction is favoured by high Po3 iri tissues. Carbonic acid ionises
into H+ and HCO;. Hydrogen ion is buffered b$ haemoglobin but aplon balance
I
is maintained by chloride shift. Reverse chloride shift occurs in the pulmonary ,
 capillaries and Carbon dioxide-is formed from carbonic acid dissociation into water
and carbon dioxide and hence carbon dioxide in gas form is exhaled.

2.9 TERMINAL OLTESTIONS

1) The passage of blood and water in fish gill is said to be countercurrent flow. What
does this mean and how does it affect gas &change efficiency? Compare the
efficiency with extraction of oxygen in the human lung.

2) Write a formula representing the reaction between haemoglobin and oxygen.

What factors influence the rate and direction of the reaction?

3) How have insects avoided the necessitybf transporting gases in the blood? Is the
system evolved by them superior to the respiratory system of mammals?

4) What happens to carbon dioxide when it enters the blood in mammals? What is
the role of carbonic anhydrase?

2.10 ANSWERS
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Self-assessment Questions
1) a) i); iii) are correct
b) knowing that oxygen percentage in air is 21
Po, = 21 x (735.18 - 18) = 150 mm Hg
100
2) Because both have high surface to volume ratio. Jelly fish also has cells lying
directly underneath the surface so that gas exchange is possible by diffusion alone.
3) When removed from the buoyancy of the water medium, the gills collapse and
tend t o stick together thus reducing the surfaces area for respiration. All
.respiratory surface have to be kept moist too, once out ~f water the gills dry up.
4) i) False ii) True iii) True iv) True v) False
 5) a) The spiracles would open.
I

b) Kerosine forms a film over water and the mosquito larvae are not able to
get air when they come to the surface.
6) iii) iv) and v)

Terminal Questions
1) Refer to sub-section 2.4.2 and sub-section 2.5.1.
respiratory membrane
2, Hb + O2 , - -- HbOz (oxyhaemoglobin).
Hint : Temperature, C 0 2 , pH and organic phosphates.
3) By evolving a tracheal system which ensures that oxygen reaches the cells directly
through a system of tubes. No system is superior. Respiratory system o f mammqls
is best adapted to their high metabolic rate and warm bloodedness. A tracheal
system is suitable for insects as it cannot supply oxygen to animals larger in size.
4) 'Refer to sub-section 2.7.3.