Gaseous Exchange

8 GASEOUS EXCHANGE

4.0 INTRODUCTION

O2 is essential for aerobic respiration.

Obtained from air, transported in blood.

CO2 is the waste product of respiration.

Transported in blood and exhaled into the
air.

Gaseous exchange (external respiration)

briefly known as the process of inhaling
oxygen from the atmosphere and exhaling
carbon dioxide to the atmosphere.

In detail, it is a physical exchange of gases

(CO2 & O2) by diffusion across the plasma
membranes/ respiratory surfaces of
organisms.

The whole structures which is responsible

for gaseous exchange is called respiratory
system.

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8.1 GASEOUS EXCHANGE IN ANIMAL

8.1.1 Types of respiratory systems in

animals

a) Respiratory Surface

Characteristics of respiratory surface to

ensure diffusion occurs at maximum,
meeting the demands of the organism
respiratory:

▪ Large Surface area

The larger the animal and the more active

it is, the larger the surface area needs for
gaseous exchange.
The greater the area exposed to the
environment, the greater the rate of
diffusion.

▪ Moist
Must be always moist so that gases can
diffuse across.

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 Thin – one cell thick ( very thin)

because diffusion is rapid over very short
distance.
e.g Simple squamous epithelium

 A good blood supply

An efficient transport will ensure that the
gases will be taken away from respiratory
surface quickly so maintaining a large
concentration gradient.

 Good ventilation gradient

A pumping system, which will continuously,
deliver gas to respiratory surfaces. In fish,
ventilation system delivers water to the
gills. In mammal, air is supplied to the lung
surfaces. This will maintain a large
diffusion gradient across the respiratory
surface. ( Biology, 7th Edition, Solomon)

Gases diffuse through the aqueous layer

covering the epithelial cells of respiratory
organs

Diffusion is a passive process/ transport.

Gasses always move from higher
concentration region to low concentration
region.

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Fick’s Law of Diffusion:

▪ The amount of oxygen or carbon dioxide
that diffuses across the membrane of an
respiratory surface depends on the
differences in partial pressure on both
sides of the membrane and also on the
surface of the membrane.

R=Ax ∆p
d

Rate of diffusion ∝ Surface area x Concentration difference

Diffusion distance

▪ Rate of diffusion (R) across a membrane

depends on:
 Surface area (A)
 Concentration (partial pressure)
difference ∆ p
 Distance (that the diffusing gases must
travel ) d

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The
The relationship between the rate of
diffusion with those factors?

 Proportional to surface area and

concentration difference but inversely
proportional to distance.
 The steeper the gradient/ the greater the
difference, the faster the rate of
diffusion.
 The larger the surface area, the greater
the rate.
 If the distance is doubled, diffusion
takes 4x longer (Solomon & Raven)

b) Types of respiratory surfaces

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LE 42-19
Respiratory
medium O2 CO2 Respiratory
(air or water) surface

Organismal
level

Circulatory system

Cellular level

Energy-rich
fuel molecules Cellular respiration ATP
from food

Respiratory structures are adapted for gases

to be exchanged in air or water.

▪ Gills adapted for respiration in water.

▪ Trachea and lungs terrestrial respiration

Advantages of gas exchange in air over gas

exchange in water:-

▪ Oxygen is more concentrated in air than

water.
▪ Oxygen diffuses more rapidly through air
than water.

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 Air is less dense and less viscous. Less

energy needed to remove air over trachea
and lungs than gills.

Disadvantage:
▪ Terrestrial organisms lose water during
respiration.

Four main types of respiratory surfaces have

evolved in animals: (Refer Figure 44.1 pg 858,
Solomon)

▪ The animal’s own body surface

( skin)..cutaneous respiration
▪ Tracheal tubes
▪ Gills
▪ Lungs

The body surface

▪ Adapted for gas exchange

▪ Small animals/Single celled organisms
respire by diffusion through the surface of
the body
▪ E.g. Annelids, shell-less mollusks, and

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amphibians.

Tracheal tube systems (Refer Figure 42.22 /pg 887)

▪ Insects and a few other arthropods

▪ Air may be brought in by actively pumping
air in through the spiracles
▪ Branching of the tracheal tubes extend
throughout the body
▪ Deliver air directly to the cells
▪ Efficient in insects because their small
bodies give a high surface area to volume
ratio.
▪ Water loss prevented by closing external
opening/spiracles.

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LE 42-22a

Air sacs Tracheae

Spiracle

LE 42-22b
Body
cell
Air
Tracheole sac

Trachea

Air Body wall

Tracheoles Mitochondria Myofibrils

2.5 µm

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Gills
 Respiratory surfaces of aquatic animals
 Evaginations of the body surface
 E.g. Bony Fishes
(Refer Figure42.21 pg 886)

▪ The gills of fishes provide a very large

respiratory surface area for
countercurrent exchange.
▪ Location: Between the buccal (mouth)
cavity and the opercula cavity

Structure of a fish gill

▪ 4 gill arches on each side of fish head

▪ Each gill arch composed of two rows of
gill filaments
▪ Each filament contains lamellae (thin
membranous plates). Within each lamella,
blood flows in opposite direction of water.
▪ Have flexible gill covers = operculum

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LE 42-21

Oxygen-poor
blood
Lamella
Oxygen-rich
Gill blood
arch
Gill Blood
arch vessel 15%
40%

%
Water 5%

70
Operculum 30%
flow

%
%

100
60

%
90
Water flow
O2
over lamellae
showing % O2 Blood flow
through capillaries
in lamellae
showing % O2
Gill
filaments
Countercurrent exchange

How do bony fishes respire?

▪ The movement of water into mouth,

through the gills and out of the fish
through operculum due to the function of
two sets of cavity:

Buccal cavity
▪ Can be opened/ closed by opening/closing
the mouth.

Opercular cavity
▪ Can be opened/ closed by the movement
of operculum

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a) Mouth opened b) Mouth closed

(Oral valve) (Oral valve)

Jaw lowered Operculum opened

Pressure ↓ Pressure ↑
Water drawn into Water forced over
buccal cavity. the gill to opercular
Operculum closed cavity to the
outside

Water movement is one way.

In gills, blood flows in an opposite direction
to the flow of water. This arrangement is
called countercurrent flow.

Maximizes oxygenation of the blood by
increasing the concentration gradient of O2
along the pathway for diffusion.

Explain the advantage of countercurrent exchange

compared with concurrent exchange.

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Countercurrent exchange 886

1. Blood low in O2 concentration enters the

back of lamella.
2. Meet with water that already had most
of its oxygen removed as it flowed
through lamella in the opposite
direction.
3. The initial O2 concentration difference
between water (15%) and blood (10%) is
not large but it is sufficient for O2 to
diffuse from water to blood.
4. As blood flows toward the front of
lamella, it meets with water that has a
higher concentration of O2. O2
continuously diffuses from water into
the blood
5. Countercurrent flow ensures that a
concentration gradient remained
between blood and water throughout the
flow.
6. This permit O2 to continue to diffuse all
along the lamellae so that blood leaving
the gills will have an O2 concentration as
high as water entering the gills.
7. Blood attains an O2 concentration of
85%.

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8. Fish gills are the most efficient of all

respiratory organs.

Concurrent exchange
 Blood and water flows in the same
direction.
 The difference in O2 concentration high
at the front of each lamella where O2
depleted blood (10%) meets O2 rich
water entering the gill (90%).
 O2 concentration in water would
decrease as that in the blood increase
 When the concentration in the two fluids
become equal, equilibrium would be
reached, net diffusion of oxygen would
stop.
 Only about 50% of the oxygen dissolved
in the water could diffuse in the blood.
(Solomon)

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Lungs

▪ Terrestrial vertebrates exchange gases

through lungs (Refer Figure 42.23 pg 888 )

▪ Lungs are invaginations from the body

surface or an internal cavity.

Primitive invertebrate, advance

vertebrate and vertebrate respiratory
system.
g-22
Types Respiratory
System
Fish  Early lobe-
finned
fishes had
lungs that
were
adaptive for
the
droughts
during the
Devonian
period
 Most
modern fish
have swim

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bladders,
which aid in
buoyancy
control

Amphibians  Many
amphibia
n
rely on
cutaneou
s
respiration
in
addition to
lungs

Reptiles  Reptilian
lungs are
relatively
simple,
with some
inner folds
to increase
surface
area

Birds  Birds have

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air sacs,
which
LE 42-25

increase
Air Air
respiratory
Anterior
air sacs
efficiency
Trachea
Posterior
air sacs Lungs Lungs (Refer
Air tubes
Figure 42.25
(parabronchi)

INHALATION EXHALATION
in lung 1 mm
pg 889)
Air sacs fill Air sacs empty; lungs fill
 Air flow is
one-way
and flow of
air is
opposite of
that of the
flow of
blood
(countercur
rent flow)
 The
parabronchi
are the site
of gas
exchange

Mammalia  Mammalian
lungs
have
millions of

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alveoli that
increase
the
surface
available
for gas
exchange
Fig 42.23 Page
888

8.1.2 Human Respiratory System

The mammalian respiratory system consists of an

airway and lungs (Fig 42.23 Page 888).

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LE 42-23

Branch
Branch from
from pulmonary
pulmonary artery
vein (oxygen-poor
(oxygen-rich blood)
blood)

Terminal
bronchiole

Nasal
cavity

Pharynx
Alveoli
Larynx
Left

50 µm
Esophagus lung
Trachea

Right
lung

50 µm
Bronchus

Bronchiole

Diaphragm
Heart SEM Colorized SEM

▪ The airway conducts air into the lungs.

 The nasal cavities filter, warm, and

moisten inhaled air
 Mucus traps particles in inspired air, and
cilia move it back towards the throat
 The nasal and oral cavities connect via the
pharynx
 The pharynx leads to the larynx, a
cartilaginous structure adapted for sound
production
 The epiglottis covers the entrance to the
larynx during swallowing
 Air passes from the larynx to the trachea,
which is supported by cartilaginous rings

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 The trachea branches into two bronchi

 The trachea and bronchi are lined with a
ciliated mucus membrane
 The two lungs are divided into lobes
 The lungs are covered by pleural
membranes, which form a pleural cavity
 The pleural fluid provides lubrication during
breathing
 Bronchi branch many times, ultimately
forming the millions of microscopic
bronchioles
 The bronchioles lead to alveoli, which are
composed of a single layer of epithelial
cells..simple squamous
 The alveoli are highly vascularized

Nasal cavity/buccal cavity

↓
Pharynx
↓
Larynx (voice box)
↓
Trachea
↓
Bronchi
↓
Bronchioles
↓
Alveoli - site of gaseous exchange

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▪ The alveoli in mammalian lungs provide a

large respiratory surface area but do not
permit counter current exchange.

8.1.3 Role of partial pressure (page 891,

Biology 7th Edi., Campbell)

▪ Is used to compare the proportions of

gases in a mixture.
▪ The partial pressure of a gas in a mixture
of gases = the pressure exerted by that
gas.
▪ Apparatus that measures air pressure =
barometer
 Barometric pressure of the air at sea level
=760 mm Hg ↔ one atmosphere of pressure
▪ Indicated by PO2 = partial pressure of O2
▪ To calculate partial pressure;

 Fractional composition of each gas in

air X atmospheric pressure
 Ex: PO2 = 21% x 760 = 160 mm Hg

8.1.4 Respiratory pigments

▪ Conjugated protein
 Consists of four polypeptide chains – two
alpha chain and two beta chain

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 Each chain combined with heme group

 Each heme group has a central iron atom,
which can bind to a molecule of O2 ∴

LE 42-28

Heme group Iron atom

O2 loaded
in lungs

O2 unloaded
in tissues

Polypeptide chain

 Haemoglobin (Hb) forms a reversible bond

with oxygen

Hb + 4 O2 ↔ HbO8 (Oxyhemoglobin)

Oxyhb releases O2 ↔ deoxyhemoglobin

(Bright red) (Dark red colour)

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 Hb = oxygen carrier protein in all

vertebrates and invertebrates ( annelids,
mollusks, echinoderm, flatworm)

▪ Hemocyanin, copper-containing respiratory

pigments = Other invertebrates (some
mollusk and artropods)
 Not in blood cell → dissolved in the
circulating fluids (hemolymph)

8.1.5 Oxygen transport and Bohr Effect

Transportation of Oxygen

▪ Plasma alone cannot supply sufficient

oxygen in a human
▪ O2 does not dissolve well in water, so that
most O2 is transported in combination with
haemoglobin.

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Oxygen-hemoglobin dissociation curve:

Oxygen saturation of hemoglobin (%) plotted
against different value of partial pressure of
oxygen (PO2) (mmHg)
▪ The curve is an S or sigmoid shape.
(Refer Figure 44-10 pg 867,Solomon &
Figure 42.29, pg 892, Campbell)

LE 42-29a
O2 saturation of hemoglobin (%)

100 O2 unloaded from

hemoglobin
during normal
80 metabolism

60
O2 reserve that can
be unloaded from
40 hemoglobin to
tissues with high
metabolism
20

0
0 20 40 60 80 100

Tissues during Tissues Lungs

exercise at rest
PO2 (mm Hg)

PO2 and hemoglobin dissociation at 37°C and pH 7.4

PO2 mm Hg
Air in alveolus 104
Blood leaving alveolus 100

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Blood leaving tissue 40

 At blood PO2 104 mm Hg, 97% of blood

Hb is saturated with oxygen.
∴ oxyHb saturation= 97%
 Blood leaving tissue (40mm Hg), oxyHb
saturation = 75%
 Amount of O2 unloaded to tissue at rest =
22% ↔ 1/5 O2 unloaded to tissue, 4/5
reserved in the blood.
LE 42-29b
O2 saturation of hemoglobin (%)

100
pH 7.4
80

Bohr shift:
60 additional O2
released from
pH 7.2 hemoglobin at
40 lower pH
(higher CO2
concentration)
20

0
0 20 40 60 80 100
PO (mm Hg)
2

pH and hemoglobin dissociation

Function of reserve:
▪ Supply of O2 during rest and exercise.
 During exercise, muscles use O2 from
capillary blood. Venous blood PO2 drop.

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 Venous blood PO2 = 20 mm Hg ↔

oxyhemoglobin saturation 35% ⇒ blood
has released more O2 ≈62% O2 is
unloaded to the tissue.
▪ Ensures supply of O2 for 5 minutes when
breathing is interrupted or the heart stops
beating.

The more curve is displaced to the left, the

more readily it loads oxygen but the less
readily it releases it/the greater the affinity
for O2

Compare the oxygen-

oxygen-hemoglobin
dissociation curve for adult Hb, fetal Hb and
mioglobin (oxygen carrier protein in skeletal
muscle)

▪ Mioglobin = respiratory protein found in

skeletal muscle. Similar to haemoglobin but
has only one polypeptide chain with a single
heme group. Each mioglobin can carry only
one oxygen molecule. The oxygenated form
of myoglobin = oxymyoglobin.

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▪ Myoglobin have a hyperbola-shaped

oxygen-hemoglobin dissociation curve and
to the left of oxygen-hemoglobin dissociation
curve for Hb showing that it has greater
affinity, can combine very readily with
oxygen but releases it only when PO2 is very
low.
▪ Oxygen-haemoglobin dissociation curve for
fetal Hb at the left of oxygen-hemoglobin
dissociation curve for Hb adult (mother).
Fetus blood get oxygen from mother through
placenta. Only possible if fetal Hb has
greater affinity towards oxygen compared to
mother’s Hb.

The Bohr Effect

▪ An increase in PCO2/Lower blood pH reduces

Hb’s affinity for O2 and cause it to release
oxygen more readily.
▪ This, therefore shifts oxygen-hemoglobin
dissociation curve to the right. This effect
is called Bohr Effect.
▪ Also caused by increase in temperature.

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 Draw oxygen-hemoglobin dissociation

curve of normal pH human blood (7.4)
and when pH is 7.6 and 7.2.
 Show the effect of temperature on
oxygen-hemoglobin dissociation curve.
Use 20°C, 37°C and 43°C.

8.1.6 Carbon dioxide transport

(Refer Figure 42.30 pg 893, Campbell)
LE 42-30
Tissue cell
CO2 transport
CO2 produced from tissues

Interstitial CO2
fluid

Blood plasma CO2 Capillary

within capillary wall

CO2
H2 O

Red Hemoglobin
H2CO3 picks up
blood Hb
Carbonic acid CO2 and H+
cell

HCO3– + H+
Bicarbonate

HCO3–
To lungs

CO2 transport
HCO3– to lungs

HCO3– + H+

Hemoglobin
H2CO3 Hb releases
CO2 and H+

H2 O
CO2

CO2

CO2

CO2
Alveolar space in lung

▪ Dissolved in plasma (7%)

▪ Bound to protein portion of Hb (23%)
CO2 + amino groups in Hb
carbaminohaemoglobin
▪ As hydrogen carbonate/ bicarbonate ions

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HCO3- (70%).
 CO2 from respiring cells diffuses into
the blood plasma and then into the
RBC/erythrocytes.
 First CO2 combines with water to form
carbonic acid (H2CO3). Reaction
catalysed by the enzyme carbonic
anhydrase.
 H2CO3 dissociates, forming protons
(H+)/ hydrogen ion and hydrogen
carbonate ions (HCO3-)
 HCO3- diffuses out of the cell into the
blood plasma and transported to the
lungs.
 Chloride ions (Cl -) diffuse inwards
from the plasma to maintain electrical
neutrality. Process = chloride shift.
 The protons left inside the cell are
mopped up by hemoglobin to form
hemoglobinic acid (HHb). (Solomon)
 This force oxyhemoglobin (HbO2) to
release oxygen.
O2 diffuses out →plasma→ tissue

▪ The reverse reactions occur in lungs so that

CO2 can be exhaled.
▪ The lower PCO2 of the air inside the alveoli
causes the carbonic anhydrase to proceed

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in the reverse direction, H2CO3 converted

into H2O and CO2.
▪ CO2 diffuses out of the red blood cells and
into the alveolus following the concentration
gradient.

8.1.7 The Control of Breathing (pg 890,

Campbell)
LE 42-26

Cerebrospinal
fluid

Pons
Breathing
control
centers Medulla
oblongata

Carotid
arteries

Aorta

Diaphragm

Rib muscles

Medulla oblongata and pons are the main

breathing control centers.

The medulla sets the basic rhythm of
breathing.

The pons controls the transitions between
inhalation and exhalation.

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Sensors in the medulla, aorta, and carotid

arteries sense the carbon dioxide and
oxygen concentration (pH) in the blood.

If there is a slight drop in pH of
cerebrospinal fluid, medulla’s control center
increases the depth and rate of breathing,
eliminating excess carbon dioxide in exhaled
air. Medulla’s control centre send impulses
to the diaphragm and rib muscles and
stimulate them to contract.

If there is a slight drop in oxygen/ O2
level is depressed ( increased pH level),
oxygen sensors in aorta and carotid arteries
send impulse to the medulla, signaling it to
increase the breathing rate.

Mechanism of Breathing

Breathing = Process by which air is taken

into the lungs (inspiration/inhalation) and
then expelled (expiration) after gaseous
exchange has taken place.

Boyle’s Law= when volume of a given
quantity of gas increased, its pressure
decreased

Air flows from region of high pressure to
region of low pressure.

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During inhalation:

 Thoracic/ chest cavity enlarged → lungs

enlarged → reduces gas pressure in the
alveoli → creates pressure gradient which
draws air into lungs.
▪ Expansion/ enlargement due to combined
movement on the rib cage (rib and
sternum) by external/ internal intercostal
muscle and diaphragm.
▪ External intercostal muscles contract,
rib cage moves up and out
 Contraction of diaphragm muscle,
diaphragm flattened, volume of thorax
increase, pressure decrease so that air
drawn in.( Antagonistically during
exhalation)

8.1.8 Lungs volumes

The volume of air inhaled and exhaled can

be measured using a spirometer.

Terms to describe the volume changes of
the lungs during breathing.

▪ The tidal volume is the amount of air moved

in or out in a normal inspiration or
expiration.

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▪ The vital capacity is the maximum tidal

volume during forced breathing.
▪ The residual capacity is the amount of air
remaining in the lungs after a normal
expiration/ even after fully forced
breathing.

 Hypoventilating= If breathing is
insufficient to maintain blood gas
measurement (PCO2 ↑ )

 Hyperventilating =If breathing is

excessive for a particular metabolic rate
( PCO2 ↓)( Solomon)

8.1.9 Respiratory diseases

Disease Respiratory failure

Asthma Constriction of smooth muscles in
the bronchiolar and bronchial wall,
excess mucus secretion and
insufficient recoil of the alveoli.
Caused by allergy and emotional
upset. Results in difficulty in
breathing.
Pneumonia Alveoli filled of fluid, caused by

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chemical, bacteria (
Streptococcus), viruses, protozoa
or fungi.
Tuberculosis Mycobacterium tuberculosis ,
water-borne bacteria causes lung
damage in variety of ways. The
infectious bacteria are normally
spread through the air by coughing
and sneezing.

Lung cancer Any inhaled irritant stimulates

( pulmonary cell to grow abnormally. Individual
carcinoma) has difficulty to breathe, chests
are pain and spitting blood.
Cigarette smokers are 20 times
the risk of non smokers to have
this disease.

8.2 GASEOUS EXCHANGE

EXCHANGE IN PLANTS

8.2.1 The structure of stomata

Stomata (stoma, sing.) are embedded within

epidermis of leaves.

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They allow CO2 exchange between the

surrounding air and the photosynthetic cells
inside the leaf, and also balance the water
requirement of plants.

A stoma consists of a pore flanked by two
guard cells ( Refer to fig. 35.17, page 725),
which regulate the opening and closing of
the pore.

8.2.2 Mechanism of stoma opening and closing

For photosynthesis, green plants need CO2

and a mean of disposing of O2

For cellular respiration, plants cell need O2
and a mean of disposing CO2

No specialized system in plants for gaseous
exchange.

▪ Reason :

 Each part of plants takes care of its own

gases exchange.
 Root, stems and leaves respire at rates
much lower than animal. Only during

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photosynthesis, large volume of gases

exchange and each leaf is well adapted.

Most of the living cells have at least part of

their surface exposed to air. The loose
packing of parenchyma cells in leaves, stem,
and roots provides an interconnecting
system of air spaces.

Gas diffuses through air several thousand

times faster than through water. Once O2
and CO2 reach the network of intercellular
air spaces, they diffuse rapidly through
them.

The exchange of gases in the leaf and
young stem occurs through pores calles
stomata.

Woody stem and mature root, through
lenticels ( Solomon)

▪ Stomata opening and closing are due to

changes in guard cell turgidity.

 Stomata must be open for

photosynthesis to occur (except in CAM
plants).
 When water flows into the guard cells,
they become swollen, and the inner

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walls bend outward, producing a pore.

 Starch is hydrolyzed to sucrose, which
increases in concentration in the guard
cells.
 Stomata close when water leaves guard
cells due to decline in concentration of
sucrose (osmotically active solute).
(Refer Figure 32-5 pg 62, Solomon )

Stomata opening and closing is triggered by

light levels

▪ Blue light stimulates this opening and

closing and is detected by yellow
pigment ( Solomon) / blue-light receptors
( Campbell) in the plasma membrane of
guard cells.
▪ Blue light activates ATP synthase.
▪ ATP synthase pumps H+ out of the guard
cells, promote the uptake of K+ into the
guard cells and causes the guard cells
become turgid.
 Stomatal closing results from the exit of
K+ from guard cells to neighbouring cells,
which leads to osmotic water loss.

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Stomata open and close in response to the

movement of H+ and K+ (potassium) across
the guard cells’ plasma membranes.

▪ K+ ions transported via voltage-activated

K+channels, then Cl- ions are taken into
guard cells through ion channels in
plasma membranes.
 K+, Cl- and malate ions accumulate in
the vacuoles of guard cells.
 Water then passes into the guard cells
by osmosis, and the cells swell.
 The opening of stomata reduced by the
outflow of K+ in guard cells during late
afternoon and at dusk ,stomata
completely closed due to the presence
of sucrose(from hydrolysis of starch).
■ Water lost from the guard cells by
osmosis, which lose their turgidity, and
thus the pore closes. (Refer Figure 32-10
pg 625, Solomon)

Other factors that affect stomata

▪ A low concentration of carbon dioxiode

opens the stomata, even in the dark.
▪ Photosynthesis reduces the

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concentration of carbon dioxide,

triggering the stomata to open.
▪ If water levels are low, stomata will
remain closed even during the day.
▪ Hormonal control (particularly abscisic
acid) is also a factor. (Refer Chap 36
Solomon).
▪ An internal biological clock and
circadian rhythms can also regulate
stomata opening and closing (circadian
cycle) (Refer Chap 32 Solomon).

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