Unit 4 Respiratory System

UNIT 4
RESPIRATORY SYSTEM

Structure
4.1 Introduction 4.4 Respiration by Lungs
Objectives Respiratory System of Amphibians
4.2 Salient Features of Respiratory Respiratory System of Reptiles
System of Vertebrates Respiratory System of Birds
4.3 Respiration by Gills Respiratory System of Mammals
General Gill Structure 4.5 Summary
Respiratory System of 4.6 Terminal Questions
Cyclostomes
4.7 Answers
Respiratory System of Fishes
Accessory Respiratory Organs in
Fishes
Respiration in Amphibian Larvae
using Gills

4.1 INTRODUCTION
In the previous unit you have learnt about the structure of various types of
digestive systems that occur in vertebrates and their role in providing nutrients
to the cells of the body. These nutrients are essential for fulfilling the energy
requirements of the animal cells. The nutrients produce energy when they
undergo oxidation during the process of respiratory metabolism. The end
products generated by oxidation are energy, water, and carbon dioxide. Thus
vertebrates consume oxygen for oxidation of nutrients present within the cells.
This oxygen needs to be replenished and the waste byproducts like heat and
carbon dioxide produced during oxidation metabolism must be removed in
order to survive. The sequence of events that result in exchange of oxygen
and carbon dioxide between an organism and its environment is known as
respiration. This is done primarily by the respiratory system. The gas
exchange between the environment and blood via the respiratory surface is
referred to as external respiration. The utilization of oxygen for oxidation of
nutrients within the cells and tissues may be termed as internal respiration.
In this unit you will study the major respiratory structures i.e. gills, and lungs
that facilitate respiration in both aquatic and terrestrial vertebrates.
109
Block 1 Comparative Anatomy of Vertebrates-I
Objectives
After studying this unit you should be able to:

 describe the structure and function of gills which form the respiratory
system of aquatic vertebrates like fishes and amphibian larvae to
breathe in water and air and explain how they are used;

 describe the structure and function of accessory respiratory organs in

fishes; and

 describe the structure and function of respiratory organs in terrestrial

vertebrates.

4.2 SALIENT FEATURES OF RESPIRATORY

SYSTEM OF VERTEBRATES
Respiration is the sequence of events that result in exchange of oxygen and
carbon dioxide between the environment and organism. External respiration
refers to gas exchange between the environment and blood via a respiratory
surface. Respiratory organs are of two types: (i) those that have respiratory
surface turned out forming an evagination called gills, (ii)those that have
respiratory surface turned in forming an invagination called lungs. Our own
lungs are a good example of such invagination.

The organs of respiration in vertebrates (the gills or lungs and in some cases
the skin) help in ventilation/breathing i.e. active movement of the respiratory
medium (water or air) across the respiratory surface.

Ventilation may be

 non directional (water/air flows past the respiratory surface in an

unpredictable pattern),

 unidirectional (water or air enters the respiratory surface at one point and
exits at another), and

 tidal (water/air moves in and out from one point only)

For the respiratory organs to function efficiently they must have:

1. A provision for renewing the supply of oxygen-containing medium,

namely, water or air that comes in contact with respiratory surface and
provision for removing carbon dioxide that is released from the respiratory
surface.

2. A large surface area which is provided with ample capillary network that
has an access to the external environment;

3. A thin and moist membrane surface which facilitates passage of gases.

Let us now discuss the structure of gills as respiratory organs of aquatic

vertebrates.
110
Unit 4 Respiratory System

4.3 RESPIRATION BY GILLS

Gills are the main respiratory organs in fishes and some aquatic amphibians.
They are composed of numerous gill filaments or gill lamellae, which are thin
walled extensions of the epithelial surface. Each gill contains a vascular
network. Blood is brought extremely close to the respiratory surface, thus
facilitating ready exchange of gases.

4.3.1 General Gill Structure

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 arches on either side (Fig.4.1a). The gill arches
separate the opercular and the buccal cavities. From each gill arch extend two
rows of gill filaments, Fig. 4.1b shows the arrangement of gill filaments in the
arches. The tips of the filaments of adjacent arches meet forming a sieve like
structure through which the water flows.

(a)

(b)

(c)

Fig. 4.1: a) Position of gill arches beneath the operculum on the left side of fish.
The operculum has been lifted to show the arch; b) Part of two
adjoining gill arches with their filaments. 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 filament showing the flat lamellae; the flow of water is
opposite to the direction in which the blood moves.

Fish gills are covered by a thin epidermal membrane which folds repeatedly to
form plate like lamellae that look like free flaps (Fig 4.1b). The structure of
lamellae increases the surface area for respiration or exchange of gases. The
flow of water is opposite to the direction in which the blood moves (Fig 4.1c).
The area of gills present in different fishes varies depending on the activity of
fishes. Fishes which are more active have larger gill areas. The rate of 111
 Block 1 Comparative Anatomy of Vertebrates-I
respiration in fishes is dependent on the amount of dissolved oxygen in water
e.g. rate of respiration is more in fishes that live in waters with low oxygen
content.

Gills are of two types : (i) external gills and (ii) internal gills.

(i) External gills (Fig.4.2) develop from the integument covering the outer
surfaces of branchial/visceral arches and protrude into surrounding
water. They are found in fish and larvae of many vertebrates including
lung fishes, amphibians etc. They are usually branched; filamentous
structures derived from ecotoderm. The functioning of external gills
poses no problem since the gill filaments are in direct contact with water
containing dissolved oxygen. The disadvantage of external gills is that
they are easily damaged.

Fig.4.2: External gills of

a salamander
larva.

Fig. 4.3: Operculum of a bony fish removed to show internal gills.

(ii) Internal gills (Fig.4.3) are associated with pharyngeal slits (series of
openings in the pharyngeal region between digestive tract and the
outside of the body) and pouches (in some vertebrates, diverticula from
the gut in the pharyngeal region that never break through to form open
passageway to the outside; such diverticula are called pharyngeal
pouches). Internal gills are composed of a series of parallel gill lamellae
although in some forms they may be filamentous. They may be borne on
both sides of the interbranchial septa but in some cases are present on
only one side. A series of gill lamellae present only on one side of an
interbranchial septum are termed as half-gill or hemibranch. Two
hemibranchs join with interbranchial septum to form a complete gill or
holobranch (Fig. 4.4).The gills are covered and protected laterally by
Fig.4.4: Hemibranch and soft skin folds such as interbrachial septum or by firm operculum. It is
holobranch. generally assumed that internal gills are derived from endoderm,
although the exact origin is not clear.

There are two main types of internal gills in fishes. The first and more
primitive types of internal gills are characteristic of elasmobranchs
(cartiligenous fishes). In this group the interbranchial septa are exceptionally
well developed and extend beyond the hemibranchs (Fig. 4.5a). The
interbranchial septa, in addition to separating the gill clefts, serve to protect
the gills themselves. The second type of internal gills is found in bony fishes.
112 In these forms, the interbranchial septa are reduced to varying degrees
Unit 4 Respiratory System
(Fig. 4.5b) from which hemibranchs protrude into a single branchial or
extrabranchial chamber located on each side between the operculum and
gills. The operculum, protects the gills in the branchial chamber which opens
to the outside through a single gill aperture. Opening and closing of operculum
bring about gill respiration in these fishes.

Fig. 4.5: Types of gill in fishes (a) elasmobranch; (b) teleost.

The gills are provided with dense capillary beds in the branchial region. Blood
is carried very close to the respiratory surface, thus facilitating ready exchange
of gases. In gills of most fishes ventilation or breathing is unidirectional. When
internal gills are used in respiration, water containing dissolved oxygen enters
through the mouth and passes through the internal gill slits into the gill clefts.
As the water passes over the gill lamellae, oxygen is taken from the water and
carbon dioxide is released. The water then passes through the external gills
slits to the outside.

Let us now study respiratory systems of different vertebrates that use gills as
their respiratory organs but before we do that try the SAQ given below.

SAQ 1
1. Indicate whether the following statements are true or false :

(a) External respiration would mean utilization of oxygen for oxidation of

nutrients in cells and tissues. T/F

(b) One of the criteria for efficient functioning of the respiratory organs is
that the respiratory surface should be thin and moist to facilitate
exchange of gases. T/F

(c) Internal gills develop from the integument covering the outer surface
of branchial arches. T/F

(d) Holobranch refers to a series of lamellae on one side of an

interbranchial septum. T/F

4.3.2 Respiratory System of Cyclostomes

Cyclostomes are jawless vertebrates, and include the hagfishes (class:Myxini)
and lampreys (class:Petromyzontida) (Fig.4.6). Among hagfishes the number 113
Block 1 Comparative Anatomy of Vertebrates-I
of gill openings on each side vary (1 to 15) with the species. In lampreys,
seven pairs of internal gill slits open from the pharynx into seven pairs of gill
clefts, which are rather large and spherical in shape.

Fig. 4.6: Placement of gill slits (pharyngeal slits) in (a) hagfish and (b) lamprey.

Respiratory medium in hagfishes is water and its flow is unidirectional. The

water enters through a single nasal opening, connecting to the pharynx via a
broad tube called the nasopharyngeal tube, enters the gill pouches and exits
through the numerous gill slits. In Myxine however there is only one gill
opening on each side. In Figure 4.7 (a) you can see the external gill openings
are located near the midventral line at some distance from the anterior end.
The gill pouches connect to the pharynx internally by backwardly running
channels that unite to form a single tube that opens on either side by a single
external branchial opening (Fig. 4.7 b and c). In other species there may be 5
to 16 gill pouches, each opening externally through separate gill slits.

Fig. 4.7: (a) Lateral view of hagfish Myxine showing the external gill opening on
one side; (b) sagittal section of head/anterior region showing
respiratory structures of hagfish; c) Diagram, showing the relation of
gill pouches to the pharynx and to the single pair of gill aperture in
114 Myxine.
Unit 4 Respiratory System
Breathing or ventilation in adult lamprey unlike hagfishes is tidal i.e. water
enters and is expelled through the gill slits. However, in larvae lamprey
ventilation is unidirectional as water is drawn into mouth and then pumped
outside from the gills in a continuous flow.

Lampreys are only one of its kind amongst living vertebrates as they have a
single nasal opening on their head which is connected to pituitary
(hypophysis) with help of a duct and ends in a blind sac called the
nasohypophysial pouch. Thus external nare does not connect to the pharynx
but ends in a blind sac The pharynx is subdivided into a dorsal oesophagus
and the ventral respiratory tube. This respiratory tube is separated from the
mouth by a flap called the velum which prevents water from flowing out of the
respiratory tube into the mouth while feeding (Fig. 4.8). Seven pairs of gill
pouches are present between the respiratory tube and body wall which open
just behind the head. The gill pouches have numerous gill lamellae developed
on their inner surface that are separated from one another by wide
membranous interbranchial septa. The gills pouches open internally by slits
into the respiratory tube and they open separately to the exterior by external
gill slits. Muscles surround gill pouches, and these together with elastic
cartilages and appropriate valves pump the water tidally in and out of the
external openings. In fact exhalation is the active movement and passive
elastic recoiling of the branchial basket results in inhalation. Adult lampreys
cannot ventilate the gills in a flow-through fashion as much of their time is
spent with their sucker like mouth affixed to bodies of other fishes. Therefore,
respiration is by tidal ventilation. However, in larval stages the water enters
through the mouth and leaves through the external gill slits.

Fig.4.8: Longitudinal section of Petromyzon showing the internal structure. Note

the division of the pharynx into a dorsal oesophagus and ventral
respiratory tube and the gill pouches that connect to it and open
externally through gill slits.

4.3.3 Respiratory System of Fishes

Ventilation or breathing in fishes is also unidirectional. Water enters the mouth
and pharynx and is expelled through external gill slits in Elasmobronchii and
through operculum in teleosts. 115
Block 1 Comparative Anatomy of Vertebrates-I
Counter current Exchange

Gas exchange takes place in gill lamellae as water flows between them in one
pO2 reflects the direction and blood within them in the other direction. This is called
amount of oxygen gas countercurrent flow. This type of flow has an important consequence. It
dissolved in the permits the fish gills to have the highest possible oxygen levels. Figure 4.9
blood. It primarily shows the advantage of countercurrent flow. Now let us suppose that the
measures the
incoming blood in gills is devoid of all oxygen, and imagine the flow of water
effectiveness of the
lungs in pulling and blood is in the same direction i.e. it is concurrent (Fig. 4.9a) when the two
oxygen into the blood streams come in contact oxygen is transferred from the water to blood at a
stream from the high rate till an equilibrium is reached after which no transfer occurs. Now
atmosphere. consider Fig. 4.9b which shows countercurrent flow. When blood which has
zero pO2 (Partial Pressure of Oxygen) comes in contact with water for the first
time, the water also has low pO2 (since it has been losing oxygen on its way to
this point) but still sufficiently more than the pO2 in blood for a pressure
gradient to be maintained and O2 flows from the water into the blood. 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.

Fig 4.9: a) Concurrent flow: When blood and water move in the same direction,
then no further exchange of O2 takes place after equal concentration is
reached. In this condition the pO2 in blood reaches the pO2 levels of the
out flowing water; b) Countercurrent flow: favours better absorption of
oxygen by blood. Blood enters with low pO 2 but leaves the lamellae with
nearly the same pO2 as water; Counter current flow also reduces the
energy cost of pumping water over the gills.

Gills arches

A series of skeletogenous gills or visceral arches encircle the pharynx of

fishes and of tetrapods. In fishes these arches primarily support the gills. They
are located between the gill clefts, one behind the other at the bases of the
interbranchial septa. The first arch is called mandibular arch and the second,
hyoid arch. The remaining visceral arches are referred to by numbers (3, 4, 5,
116 6, etc.).
Unit 4 Respiratory System
The first gill pouch or cleft lies between mandibular and hyoid arches and is
often referred to as the hyomandibular cleft. In fishes it is either modified to
form a spiracle or is closed altogether. The arrangement of gill arches and gills
in elasmobranchs and bony fishes is shown in Fig 4.10.

Fig. 4.10: Arrangement of gill arches and gills in (a) an elasmorbranch and (b) a
teleost. Notice that bony fishes have a highly reduced interbranchial
septum.

In Acipenser, Polydon, Polypterus, the first gill pouch has become modified
and opens to the outside by means of a spiracle. Rudimentary gill lamellae
may be located on the anterior wall of the spiracle. Since blood supply to
these lamellae consists of oxygenated blood, they do not perform a respiratory
function and the term false gill or pseudobranch is applied to them. The
spiracles generally open on the top of the head and in some species they are
provided with valves.

(I) RESPIRATION IN CARTILIGENOUS FISHES

Living cartiligenous fishes are placed in class Chondricthyes which includes

mostly marine sharks, rays and skates (Elasmobranchii); and elephant
fishes, rat fishes and rabbit fishes (Holocepahli). In Elasmobronchii five to
seven gill arches and gill slits in separate clefts on either side along pharynx
are present. Elasmobranchii lack operculum. Holocephali has 4 gill slits
covered by cartilaginous operculum.

In Elasmobronchii breathing/ventilation occurs when large volume of water is

sucked into the buccal cavity via the mouth and spiracles. The expansion of
buccal cavity causes an increase in volume of water sucked in. The mouth
and spiracle close and muscles around the buccal cavity contract forcing
water past the gills and out from the external gill slits (Figure 4.11). 117
Block 1 Comparative Anatomy of Vertebrates-I

Fig. 4.11: Water ventilation in Elasmobronchii. Blood flow is countercurrent to

the blood flow.

Most elasmobranchs are pentachid i.e. have five pairs of clefts in addition to
the spiracles. Another form of shark, Hexanchus, has six and another
Heptanchushas seven clefts. Heptanchus also has the largest number of gill
clefts of any gnathostome (jawed vertebrates).

(II) RESPIRATION IN BONY FISHES

Recall from the previous course (BZYCT-131),that bony fishes includes the
ray finned fishes placed in Class Actinopterygii which has subclasses (i)
Chondrostei where spiracles are present; eg. Polydon, Acipenser (ii)
Neopterygii: (gars, bowfins and teleosts); includes Amia, Lepistoseus and
most of the world’s bony fishes and (iii) Cladistia where spiracles and lungs
are present eg., Polypterus. The other group of bony fishes includes the lobe-
finned fishes placed in Class Sarcopterygii. These fishes are fishes with
lungs.

The gill apparatus of bony fishes have five gill slits, a bony operculum which
arise from the hyoid arch and moves backward over the gill chamber.
Operculum provides a protective cover over the branchial arches and gills they
support (Fig. 4.12a) . In bony fishes breathing or ventilation occurs when water
enters buccal cavity through mouth over the gills as simultaneously the
opercular cavity expands. Expansion of opercular cavity causes decrease in
118 pressure so water enters opercular cavity. When opercular valve opens water
Unit 4 Respiratory System
flows out of the opercular cavity. Water is driven across the gills by two
pumps; buccal pump and opercular suction. Opercular movement assists in
the expulsion of water from gill chambers (Fig. 4.12b).

Fig.4.12: a) Teleost fish (lateral view and horizontal section) b) ventilation cycle
in teleosts.

Most actinopterygian fishes have structures called swim bladder or air bladder
between alimentary canal and vertebral column. The distended sac like air
bladder varies in length and form in different fishes. It is essentially hydrostatic
in function and aids in boyancy of fishes (see Box 4.1). 119
Block 1 Comparative Anatomy of Vertebrates-I
Box 4.1: SWIM BLADDER

All fishes are slightly heavier than water. This is because the skeleton and the tissue
of fishes contain certain heavy elements that are present only in trace amounts in
water. The bony fishes are aided in buoyancy in water by having a floatation device—
a gas filled space—the swim bladder. Swim bladder is present in most pelagic fishes
but is absent in fishes that live in abyssal waters and bottoms of the sea. The neutral
buoyancy is achieved by adjusting the volume of the gas in swim bladder. Such an
adjustment would enable the fish to remain suspended indefinitely at any depth with
no muscular effort. Gas is added to the bladder when the fish descends to greater
depth, and when it swims up, gas is removed from the bladder making the fish lighter.
Swim bladders differ from lungs in three ways. First, swim bladders are usually
situated dorsal to the digestive tract, whereas lungs are ventral. Second, swim
bladders are single, whereas lungs are usually paired. Third, in swim bladders,
returning blood drains to the general systemic circulation (cardinal veins) before
entering the heart. In lungs, venous return enters the heart separately from the
general systemic circulation

Lung fishes are air breathing forms and include; Polypterus, Lepidosiren,
Neocertodus and Protopterus. The lungs fishes are characterised by either a
bilobed lung as in most cases or just one lobe as in Neoceratodus. The
presence of external gills is rare among lung fishes. In Polypterus a single pair
of external integumentary gills is present in the region of the hyoid arch and a
bilobed lung developed symmetrically and the duct opens ventrally into
pharynx (Fig. 4.13).

Fig. 4.13: The bilobed lung of Polypterus.

The epithelial lining of the lung is not smooth and there are a few furrows that
increase the surface which is in contact with air. In Polypterus, lungs are
supplied by pulmonary arteries off the sixth embryonic aortic arch, but the
venous return is through the hepatic veins. Neoceratodus comes to the
surface to breathe air but only when oxygen tension falls below 83 mm Hg and
is known to survive foul waters that kills other fishes, though, the fish cannot
live out of water. The other lungs fishes, Protopterus and Lepidosiren, have
been shown to obtain nearly 98% of their oxygen requirements from air. The
floor of the mouth is lowered and air drawn in through the mouth is forced
back by a buccal pump, with the mouth being closed and the tongue pressed
against the roof as a seal. The hyoid apparatus and the pectoral girdle help in
the process of inspiration. Expiration is exclusively by the elasticity of the
lungs, with ribs playing no role in the process. The gills in most lungfishes
except Neocerotodus are atrophied and do not allow adequate air exchange

Most fishes die soon after being exposed to air, even though their gills are
kept moist. Lack of water in the branchial chambers as well as the
accumulation of mucus causes the gills to stick to each other. With the result
120 the exposed respiratory surface is decreased and the exchange of gases is no
Unit 4 Respiratory System
longer adequate. Fresh-water fishes face the problem of their environment
getting dried up and to overcome this problem, in addition to gills, they have
evolved accessory organs for breathing air. In the following subsection you will
briefly learn about the accessory respiratory structures of certain fresh water
fishes.

Before you proceed to the next section attempt the given SAQ to test your
understanding of what you have learnt about respiration in fishes.

SAQ 2
1. Fill in the blanks with suitable words.

(a) The number of pairs of internal gill slits present in lampreys is

………………. .

(b) In lampreys inhalation is a ……………… process and exhalation is

an ……… process.

(c) The first visceral arch of vertebrates is ……………… arch and the
second is………………

(d) The system of water flow across the gills in fishes that ensures an
80% oxygen uptaken is knows as ………………… ……………. flow.

(e) In elasmobranchs …………………… ………………. are well

developed and extend beyond hemibranchs.

2. Pick out the false statements and correct them.

(a) Interbranchial septum support gill filaments in both bony and

cartilaginous fishes.

(b) Spiracles are present in all elasmobranchs.

(c) All bony fish have only one external gill slit that opens to the exterior
on each side.

(d) All elasmobranchs lack opercula.

4.3.4 Accessory Respiratory Organs in Fishes

The accessory respiratory organs of fishes are the outgrowths either of the
pharynx or the branchial chamber that are richly supplied with blood vessels.
In most air-breathing fish, wide gill filament and secondary lamellar spacing
prevents coherence and collapse. Air from outside is drawn into these
chambers by mouth and retained there for aeration of the blood.
Actinopterygians like Anabas, Ophiocephalus, Amphipnous, Clarias, and
Saccorbranchus are some of the fishes provided with accessory respiratory
organs.

Anabas, the climbing perch, migrates from pond to pond and during such
times while it is on land, breathes air from the atmosphere. The two air
chambers are the extensions of the branchial cavities and lie on each side of 121
Block 1 Comparative Anatomy of Vertebrates-I
the head. The accessory organs, also known as labyrinthine organs (Fig.
4.14) are outgrowths of the upper part of the first branchial arch. Each organ
consists of concentrically arranged wavy plates, covered by a vascular
membrane. The air chamber communicates with pharynx by an opening
situated between hyoid and the first branchial arch. Essentially air that is
drawn by the mouth passes into the branchial chamber and exits through the
branchial aperture. Anabas can remain outside water for nearly six to seven
hours. It acquires about 54% of its oxygen requirement from air .

Fig. 4.14: The labyrinthine organ of Anabas.

Ophiocephalus, the murrel, also has a pair of air chambers one on each side
of the head (Fig. 4.15). Each air chamber arises as an outgrowth of pharynx
above the first gill arch and extends as far as the last gill cleft. Air enters into
the chamber by the mouth and leaves through the opercular arch.

Fig. 4.15: The accessory respiratory organ of Ophiocephalus.

The Indian catfish Clarias, has the most complicated and highly branched
vascularised paired accessory respiratory organs. These consist of a
suprabranchial chamber with a highly vascularised membrane and
branching arboriform or dendriform organs (Fig. 4.16) lying in
suprabranchial chamber. These are more specifically derived from the upper
122 parts of the second and fourth branchial arches. They are tree like supported
Unit 4 Respiratory System
by cartilaginous internal skeleton and the ends of each branch has a knob of
cartilage core covered by a highly vascular membrane. There is a well
developed inhalant and exhalent aperture in the suprabranchial chamber. The
fish comes to the surface to take in gulps of air which goes in the
suprabranchial chamber via the pharynx. The air is exhaled via the exhalent
aperture by the contraction of the chamber.

Fig. 4.16: The arboriform accessory respiratory organ of Clarias.

In Amphinous the air chambers (Fig. 4.17) arise as saccular outgrowths of

dorsal wall of the pharynx extending as far as the third branchial arch. The
walls of the sacs are folded and vascular. The sacs communicate with pharynx
by an opening through which air is drawn in. The air exits through the gill slits
and opercular opening. The gill filaments of the first gill arches are highly
reduced.

Fig. 4.17: The air chamber in Amphipinous. 123

Block 1 Comparative Anatomy of Vertebrates-I
In Saccobranchus there is a pair of tubular sacs that arise as outgrowths of gill
chambers extending upto the middle of the tail region (Fig. 4.18). The folds in
these tubes form a sort of air chamber that communicates with the buccal
cavity by a slit. The air passes in an out of the chamber through the slit. The
mudskipper Periophthalmus that lives in brackish waters has large opercular
cavities that are filled with air drawn through the mouth. Mudskipper is so
accustomed to life out of water; it dies of suffocation if prevented from living on
land for a long period. A similar arrangement in which the opercular cavity
functions as an aerial respiratory organ is noticed in the tropical rock skipper
(Andamia).

Fig. 4.18: The accessory respiratory organ of Saccobranchus.

4.3.5 Respiration in Amphibian Larvae using Gills

Most amphibians spend their larval life in water and, after metamorphosis into
adults move to the land. During larval life, external gills are used as organs of
gas exchange. In addition the moist and highly vascularised skin also serves
as the respiratory organ. Skin and buccal cavity are the routes of oxygen entry
for respiration. The amphibians respire through skin, external gills, internal
gills and the lungs.

The three pairs of external gills that appear at the time of hatching are
branched processes connected with the blood vessel from aortic arches.
Subsequently folds from the posterior edges of the hyoid arches cover the gills
as operculum. At a later stage the gills are resorbed.

The internal gills are formed with the establishment of mouth opening. The
internal gills consisting of branchial filaments are formed ventral to the external
gills on the branchial arches and project into the opercular cavity. The third,
fourth and the fifth visceral arches possess two rows of filaments and the sixth
arch has only one row on its anterior side. Blood vessels from aortic arches
provide vascular connections. Water passes from mouth to pharynx and then
through the gill slits into the opercular chamber. It finally leaves through the
124 opercular aperture. The internal gills disappear at the time of metamorphosis.
Unit 4 Respiratory System
In a few urodeles, gills are retained thoughout life, but in most urodeles and all
the tailless amphibians, they disappear at the time of metamorphosis. Newly
developed lungs then take over the function of respiration.

Structures analogous to gills may develop in reptiles, birds and mammals. In

reptiles five pairs of pharyngeal pouches are formed during embryonic life and
in birds and mammals only four develop. In the latter groups a fifth one may
also develop, but it remains rudimentary and attached to the fourth pair. The
pouches do not break through to the outside, but very occasionally, they may
do so. If the pharyngeal pouches fail to disappear in the normal manner, they
may lead to the formation of branchial cysts and fistulae.

SAQ 3
1. Match the type of accessory respiratory organ with the fish that possesses
it.

(a) Air Chamber i. Amphipnous

(b) Tubular Chamber ii. Anabas

(c) Arboriform organ iii. Ophiocephalus

(d) Saccular Chamber iv. Saccobranchus

(e) Labyrinthine organ v. Clarias

2. Choose the correct answer from the alternatives provided.

(a) The internal gills appear appear/disappear at the time of

metamorphosis.

(b) In a few tailless amphibians/urodeles, gills persist throughout adult

life.

(c) Gills develop/do not develop in association with the pharyngeal

pouches of reptiles, birds and mammals.

(d) If the pharyngeal pouches fail to disappear/appear, they may lead to

the formation of branchial cysts and fistulae.

4.4 RESPIRATION BY LUNGS

One of the most important changes in vertebrate evolution was the transition
from water breathing to air breathing. In terrestrial animals the main
respiratory organ is the lung. Vertebrate lungs are elastic bags designed for air
breathing (Fig 4.19). The volume of lungs expands when air is inhaled and
shrinks/contracts when air is exhaled. Embryologically, lungs develop from
outpocketing of endoderm from the pharynx. The diverticulum divides into two
halves, the lung buds, which are destined to give rise to the bronchi, and the
lungs proper. The original unpaired duct, which connects the lungs to the
pharynx, serves to carry air back and forth and is known, in most cases, as
windpipe or trachea. Generally, the trachea branches into two bronchi, one to 125
Block 1 Comparative Anatomy of Vertebrates-I
each lung. In some species, each bronchus branches into successively
smaller bronchioles that eventually supply air to the respiratory surfaces within
the lung. In snakes which have slender bodies one lung may be reduced in
size; or may be absent.

The trachea and bronchi bring the air we breathe to the lungs. Each bronchus
is shaped like a tree, with lots of smaller and smaller branches. They may
branch to varying degrees, depending upon the species. The smallest
branches are called bronchioles and at the end of these are air sacs (alveoli).
They are about 600 million alveoli (large surface area provided with ample
capillary network) in the lungs in humans. These are covered with capillaries,
and here the exchange of gases takes place.

While evaginated gas exchangers like gills can be ventilated continuously and
unidirectionally through buccopharyngeal pump or ram ventilation, the lungs
developing as invaginated organs, and having a narrow entry/exit point to
atmospheric air, can only be ventilated tidally, i.e. bidirectionally (= in-and-
out). As a result, dead space is created in the major air-conducting passages.
The trachea, bronchi, bronchioles and alveoli holds a significant volume of air.
During exhalation most of the air from the lungs is expelled, however some air
is left in the passageways. On inhalation, this air is again drawn into the lungs.
This volume of air within the respiratory passageways is called the dead
space. The total volume of air that is inhaled in a single breath/inspiration is
known as the tidal volume. Normal tidal volume of humans at rest is
approximately 500 ml. As the dead space is about 150 ml (30%), only 350 ml
(500 ml—150 ml) of fresh air actually reaches the lungs. Lets us now see the
structure of lungs and the respiratory system in different groups of vertebrates
and how they have evolved/adapted to their environment.

126 Fig 4.19: Lungs in terrestrial vertebrates.

Unit 4 Respiratory System
4.4.1 Respiratory System in Amphibians
The amphibians are tetrapods with moist scaleless skins and include anurans
(frogs), urodeles (salamanders) and caecilians. You have read in the earlier
section how amphibians have transient internal and external gills in early A few urodeles do not
stages of their life. Adult amphibians generally respire by lungs, bucco- develop a lung bud
pharyngeal cavity as well as by skin. Respiration by skin is known as the and lack both gills and
cutaneous respiration and by lungs is known as pulmonary respiration. The lungs bud throughout
balance between cutaneous and pulmonary respiration varies among species life. A specially
vascularised region of
and within a species it depends on body temperature and animal’s rate of
the pharyngeal and
activity. Amphibians show more dependence on lungs (pulmonary) for oxygen oesophageal lining is
uptake as temperature and activity increases. used as the
respiratory membrane
(i) Cutaneous respiration along with the skin.
Salamanders
Amphibians depend on cutaneous respiration for a significant part of their gas
inhabiting swift
exchange. When submerged in water they respire using the skin which is mountain streams
membranous containing large network of capillaries. The skin of the may only have
amphibians has mucus glands distributed over entire body surface which vestigial lungs which
secrete glycopeptides and glycerol that keeps the skin moist. The skin is are few millimeters in
supplied with the capillaries of the cutaneous artery that carries the length, as buoyancy
deoxygenated blood. The respiratory exchange of gases takes place between would be a
disadvantage in
the atmospheric oxygen and the carbon dioxide carried in the blood. The
swiftly flowing
oxygenated blood is returned to the heart by the cutaneous vein and thence to currents.
the general circulation.

(ii) Pulmonary respiration

The respiratory tract of frog includes the external nostrils, nasal chambers,
internal nostrils, bucco-pharyngeal cavity, glottis, laryngo-tracheal chamber
and a pair of bronchi. The median slit-like glottis on the floor of pharynx opens
into larynx (laryngo-tracheal chamber). Lungs of amphibians appear as two
simple sacs, elongated in urodeles and bulbous in anurans. They occupy the
pleuriperitoneal cavity along with other viscera. The left lung is usually longer
except in caecilians, in which it is rudimentary. The internal lining of the
amphibian lung may be entirely smooth or have internal folds or septa that
increase the inner surface to form many chambers that increase the surface
are for respiration (Fig. 4.20).

Fig. 4.20: The Respiratory organs of frog (dorsal view). Right lung partly cut to
show inner partitions and alveoli. 127
Block 1 Comparative Anatomy of Vertebrates-I
There are two processes involved in pulmonary respiration. First is the
drawing in of the air into lungs - the inspiration and second, the forcing out or
air from lungs-the expiration. Since ambhibians do not have ribs or
diaphragm, inspiration occurs in two stages (Fig.4.21). (i) Essentially in the
first stage the air is drawn into the buccal cavity from outside. The lowering of
the floor of the buccal cavity causes an increase in buccal space. This in turn
leads to the rushing of air from outside into buccal space through external
nostrils. The buccal cavity thus functions as a suction pump. It is believed that
some gaseous exchange occurs in the buccal cavity (Buccopharyngeal
respiration). This is followed by opening of glottis which causes release of air
from the lungs into the buccal cavity. This released air mixes with the fresh
inhaled air in the buccal cavity. (ii) In the second process, the nostrils as well
as the mouth close. The floor of the buccal cavity is raised which causes an
increase in pressure; but the pressure is insufficient to open the mouth. The
air enters the laryngo-tracheal chamber through the glottis. The elastic wall of
the lungs causes the dilation of lungs allowing the entry of air and exchange of
gases takes place in the alveoli of the lungs. During expiration, the elastic wall
of the lungs recoil expelling the air contained in them. The air arrives in the
buccal cavity and from there moves outside through the nostrils.

Fig. 4.21: Stages in inspiration of frog (a) first stage (b) expiration (c) second
stage.

In Urodela (newts) where the animals are mostly aquatic, the lungs are poorly
vascularised and internal surface is smooth while in metabolically active Hyla
(tree frog) the lungs are more elaborate with many more blood capillary
meshes per cm2 than the aquatic newt. Thus we see that as animals became
primarily terrestrial their respiratory surfaces also became more suitable for
their life styles.

4.4.2 Respiratory System in Reptiles

Reptilians were the first vertebrates to adapt adequately to a completely
terrestrial life. Their dry scaly skin reduces cutaneous respiration to a
negligible level. As a result reptiles depend almost exclusively on lungs for gas
exchange, supplemented in some of the aquatic turtles by pharyngeal
membrane respiration. A larynx is present though vocal chords are absent.
The respiratory passage is supported by cartilages. Lungs of most reptile
unlike the simple sac like lungs of amphibians are composed of many small
compartments and thus have a sponge like texture and a larger respiratory
128 surface (See fig 4.19 again). This type of lung is needed because of the
Unit 4 Respiratory System
general increase in metabolic activity of reptiles. The lungs of many snakes
and most lizards typically include a single central air chamber. The respiratory
surface within the lungs is septal meaning that partitions form and subdivide
to increase the surface area exposed to incoming air. The interconnecting
septa divide the lumen into faveoli which are air compartments that open into
a central chamber within each lung. Faveoli differ from alveoli of mammalian
lungs as they are not found at the end of a highly branched tracheal system.
The thin walls of faveoli have capillary beds and are subdivided into internal
septa.

Lungs of reptiles are similar to those of mammals as they are aspiratory

(drawing in air) i.e. air is sucked into lungs by changing the size and pressure
muscle as in amphibians. Lungs change their shape and induce airflow in or
out. The reptiles fill their lungs by expanding the rib cage. The expansion
reduces the pressure in the lungs and air is drawn into lungs. Air is retained in
lungs during a period of apnea (cessation of breathing) and is then forced out
of the lungs by contraction of trunk muscles and elastic recoil of lungs.

The left lung in Amphisbaenas (limbless lizards) and in derived snakes

(Colubridae,Viperidae, Elapidae) is rudimentary or absent. The more
primitive snakes such as boas and pythons have both lungs. The elongated
lungs of snakes and limbless lizards are divided functionally into two zones,
the anterior vasculated zone and the posterior avascular, saccular zone which
stores air. In marine reptiles lungs are multi-chambered. In higher lizards,
crocodilians and turtles, the septa are so constructed that there are numerous
large chambers, each with a multitude of individual sub-chambers.

Volume of lungs is relatively larger than in mammals but the surface area is
sometimes 100 times smaller in proportion to body weight. In puffing adder
an enormous diverticulum of the left lung extends into the neck region.
Inflation of the diverticulum causes the neck to spread characteristically, and
inflation of the lungs causes the body to swell. In the spotted king snake, the
lung and its bronchus extend fully two-thirds the length of the body. The
purpose of the large volume is to provide a reservoir of air, useful in diving
species for holding breath. In aquatic forms lungs are often provided with
smooth avascularised air sacs. They are also useful for maintaining buoyancy.

The oxygen requirement of reptiles is relatively low. Their standard metabolic

rate is only 10 to 20 percent of that in homeotherms. Most reptiles are
therefore incapable of sustained activity. Their movements are in short bursts
during which their muscles contract anaerobically. However, reptiles can
tolerate much greater changes in the circulatory components of the blood than
in mammals. Such a property enables them to exist for a long period in low
oxygen conditions. Lizards, snakes and crocodiles can survive for 30 minutes
in pure nitrogen and turtles for several hours.

Fig. 4.22: The lungs of lizard. 129

Block 1 Comparative Anatomy of Vertebrates-I
Generally the respiratory system of lizards has the following features. The
glottis leads into larynx, which is supported by cricoids and arytenoids
cartilages. From the larynx the trachea passes backwards on the ventral side
of the neck. Rings of cartilage support the long trachea. It bifurcates behind
into two bronchi and each bronchus enters into a lung. The inner lining of the
lung is raised into a network of delicate ridges so as to produce a spongy or
honeycomb-like appearance. The ribs are pulled backwards and forwards by
the muscles, which extend between them, thereby altering the size of the body
cavity. When the body cavity is increased in size, air from outside passes
through the nostrils into the lungs and dilates them, resulting in inspiration.
The expiration is a passive act.

Crocodiles have a muscular diaphragm that pulls the pubis back (crocodiles
have a mobile pubis bone) which pulls the liver down, thus creating space for
the lungs to expand. This is known as the hepatic piston method of
ventilation. They have a unidirectional flow of air during inspiration and
expiration. Turtles have a rigid carapace that does not allow the type of
expansion and contraction seen in other amniotes. They use their fore-limbs
and pectoral girdle to force air in and out.

SAQ 4
1. Indicate weather of the following statements are true or false.

(a) In terrestrial vertebrates, a diverticulum that grows out ventrally from

the floor of the pharynx posteriorly to the last gill pouch develops
into lungs T/F

(b) In dipnoans lungs have only a hydrostatic function. T/F

(c) Protopterus and Lepidosiren, the lung fishes can obtain nearly 98%
of their oxygen from the air. T/F

(d) In amphibians cutaneous artery carries deoxygenated blood and the

cutaneous vein the oxygenated blood. T/F

(e) During pulmornary respiration, the buccal cavity of frogs function as

suction pump. T/F

(f) Higher reptiles have voluminous lung but with a surface area that is
100 times smaller in proportion to body weight. T/F

(g) In aquatic reptiles, lungs are provided with vascularised sacs that
have hydrostatic function. T/F

(h) Lizards, snakes and crocodiles have a higher standard metabolic

rate and hence their oxygen requirements are quite high. T/F

4.4.3 Respiratory System in Birds

Though the respiration in birds, is pulmonary as it is in reptiles, the respiratory
system of birds differs radically from that of reptiles and mammals and is
130
Unit 4 Respiratory System

marvelously adapted for meeting high metabolic requirements of flight. Fig.

4.23 shows the respiratory system of birds. The respiratory system of birds
occupies as much as 20% of the volume of the body, as against only 5% in
humans. In birds the lungs and the respiratory passages are highly modified.
The highlights of such modification are: (i) the formation of extensive
diverticula or air sacs in lungs that invade most parts of the body; (ii) the
anastomosing of the air ducts within the lungs so that no passage terminates
blindly within the lungs and (iii) isolation of lungs in pleural cavities. The air
sacs are blind thin walled, distensible diverticula of the lungs that invade most
parts of the body and store air.

(a) (b)

Fig. 4.23: The respiratory system of a bird showing: (a) location within the body;
(b) its detailed structures

The paired nostrils of birds lead to internal nares above the mouth cavity. The
slit like glottis in the floor of the pharynx opens into the long, flexible trachea.
The trachea continues to the syrinx. The voice box of birds is called syrinx.
From the syrinx primary bronchii enter the lungs and are then called
mesobronchus. The lungs are small paired, spongy organs more rigid with
little elasticity (Refer to Figs. 4.23 & 4.24).

Branching off from the mesobronchus are smaller tubes called dorsobronchus.
The dorsobronchus, in turn, lead into the still smaller parabronchus. Unlike
mammals where the tertiary branches end in sac like alveoli, the tertiary
branches, in birds are tube like also known as air capillaries, through which air
flows continuously. Gas exchange actually occurs in the air capillaries. A
series of valves ensure that flow of air remains one way.
There are nine interconnected aspiratory air sacs (Fig. 4.23) which lie in
various organs of the bird. The cervical, anterior thoracic, posterior thoracic, 131
Block 1 Comparative Anatomy of Vertebrates-I
and abdominal sacs are paired and the interclavicular sac is unpaired. The
cervical air sac lies at the base of the neck. The posterior thoracic air sac, is
close to the side wall of the body. The anterior thoracic sac is just in front of
posterior thoracic air sac. The unpaired intraclavicular sac gives off a
diverticulum or axillary air sac.The abdominal air sacs lie one on each side of
the body among the coils of the intestine. All these nine sacs extend into the
bones, forming the pneumatic cavities and replacing a substantial amount of
bone marrow in them. Some birds can thus respire through the humerus and
other bones if they are fractured and exposed even if their trachea is blocked.

The lungs of birds are capable of little expansion, as they are attached to the
ribs and thoracic vertebrae and are comparatively smaller than those of
mammals. The lungs of birds are more efficient because air flows through
them in one direction rather than back and forth (Fig. 4.24a). One way flow of
air through the lungs leads to greater concentration of oxygen at the epithelial
exchange surfaces than in other terrestrial vertebrates that ventilate their
lungs bi-directionally. Birds can thus obtain enough oxygen even when flying
at high altitude where partial pressure of oxygen is low.

Air in the bird respiratory system passes through in two breath cycles as
explained below (Fig.4.24 a and b):

(i) During the process of respiration, air which is first inhaled by the bird
passes through straight ducts namely, the bronchi and mesobronchi into
the posterior air sacs.

(ii) Inhalation of air is followed by exhalation which causes the previously

inhaled air to move from the posterior air sacs into the parabronchi of the
lungs.

(iii) The second inhalation moves this air which is present in the parabronchi
of the lungs into the anterior air sacs. The second exhalation then
passes the air through the bronchi and out of the system.

Oxygen exchange occurs both during inhalation and exhalation in this type of
respiration. The posterior abdominal and the anterior air sacs expand during
inhalation. All birds have this one-way air flow system and some may also
have a two-way flow system which is not discussed in this unit.

The raising of the sternum when the animal is at rest and lowering of the
backbone when the animal is in flight decreases the size of the body cavity,
leading to the air being forced outside (exhalation). Thus expiration is an
active process in birds. Inhalation is a process, brought about by the
rebounding of the muscles to their original size, causing an increase in the
size of the body cavity.

The lungs of small passerine birds that have a higher metabolic rate have
more efficient lungs in comparison to gliding birds that use less energy during
132 flight. Birds like fowls and the emu that do not fly much have a lower
Unit 4 Respiratory System

pulmonary blood diffusion capacity and Humboldt penguin has a much thicker
blood-gas exchange barrier in the lungs which protects the lung from
collapsing during diving due to hydrostatic pressure. Let us now see the
adaptations in the respiratory system of mammalian vertebrates.

(a)

(b)

Fig. 4.24: a) Diagram showing the pathway of unidirectional flow of air in birds.
b) Diagram showing the exhalation and inhalation in birds.

4.4.4 Respiratory System in Mammals

Mammals have a pair of lungs enclosed in a thoracic cavity. The bony
framework of the thoracic cavity is formed of thoracic vertebrae, ribs and
sternum. The lungs of the mammals are multi-chambered and usually divided
into lobes. Usually the right side has more lobes than the left side. Humans
have three right and two left lobes.

Figure 4.25 shows the respiratory organs of humans. The air from outside
enters through the external nostrils and nasal passages into pharynx. From 133
Block 1 Comparative Anatomy of Vertebrates-I
the pharynx it passes through the glottis into trachea. The trachea is a long
tube that traverses the neck and lies ventral to gullet. The anterior part of the
trachea is enlarged to form the voice box or larynx. The vocal chords are
located inside the larynx and the vibrations of the vocal chords results in the
production of the sound. The trachea bifurcates into two primary bronchi. Each
primary bronchus enters into lungs and branches into secondary and tertiary
bronchi, and finally into bronchioles. Terminal bronchioles lead into thin walled
delicate alveolar ducts, the walls of which are evaginated to form clusters of
alveoli. The lungs are protected and cushioned by the pleura. The pleura is
made of two thin layers of tissue: a) the inner layer (visceral pleura) which
wraps around the lungs and is stuck so tightly to the lungs that it cannot be
peeled off, and; b) the outer layer (parietal pleura) which lines the inside of the
chest wall. The pleura prevent the lungs from separating from the rib cage.
The very thin space between the layers is called the pleural cavity. A liquid,
called pleural fluid, lubricates the pleural cavity so that the two layers of pleural
tissue can slide against each other as the lungs inflate and deflate during
respiration.

Fig. 4.25: The respiratory system of humans

In mammals buccal cavity plays no role in respiration, and the diaphragm and
ribs play an important part. You can see in Figure 4.26, that during inhalation
the rib muscles (external intercostal muscles extend between the ribs) and
diaphragm contracts causing the raising of the ribs and flattening of the
diaphragm increasing the size of the thoracic cavity. The pressure decreases
and the air enters into lungs. The entire process constitutes inspiration.
Expiration is a passive process, brought about by the relaxation of the
intercostals muscles and the diaphragm. The thoracic cavity is brought to its
normal size and as a result the air is forced out.

The alveoli are separated by thin wall vascularised, septa that measure 50μm
in shrews and bats as they have a high metabolic rate and is as much as 1mm
in sirenians (manatees and dugongs) that are more placid. The alveolar
134 surface is covered by certain, Type I alveolar cells where gas exchange
Unit 4 Respiratory System
occurs and Type II alveolar cells that secrete the surfactant a substance that
reduces the surface tension at the air-alveolar surface so that the alveoli can
expand.

Fig. 4.26: Process of Inhalation and exhalation.

SAQ 5
1. Fill in the blanks with suitable words.

(a) In human right lungs have …………………. lobes and the left lung
has …………………. lobes.

(b) Birds have …………………. number of air sacs.

(c) In mammals inspiration is an …………… process and expiration is a

……………process.

(d) Normal tidal volume of human at rest is approximately …………. ml.

(e) The voice box of birds is called ………………….. .

4.5 SUMMARY
 The respiratory system is designed for exchange of oxygen and carbon
dioxide between the organisms and the environment.

 In all chordate embryos a series of visceral pouches develop on either

side of the pharynx. In fishes and larval amphibians perforations occur in
the pouches, forming gill slits which connect the pharynx to the outside.
The pouches are then called gill clefts. Gills are evaginations of the
respiratory surface found in aquatic vertebrates. Internal gills, supported 135
Block 1 Comparative Anatomy of Vertebrates-I
by visceral arches, are vascular, lamellar or even filamentous extensions
of the epithelial surface of the gill pouches. Internal gills are present in
cyclostomes, fishes and some aquatic amphibians. Respiratory
efficiency is increased by the countercurrent flow of blood and water in
the gills.

 External gills, generally covered with ectoderm, may project from the
outer surfaces of the visceral arches as filamentous outgrowths. They
are found in larval amphibians, are usually confined to larval life and
disappear at the time of metamorphosis. In a few urodeles, however gills
occur throughout the adult life. Of the five pairs of pharyngeal pouches
that develop during the embryonic life of amphibians, only the 2nd, 3rd
and 4th pairs are perforated and connected to the outside. Also during
metamorphosis initially external gills appear which degenerate and a
new set of internal gills develop from the tissue covering the same
visceral arches.

 A few fresh water fishes have evolved accessory respiratory structures

that are useful at times when there is an oxygen deficiency in the
medium in which they live. The highly vascularised structures are
essentially outgrowths of pharynx or the branchial chamber. Anabas,
Amphipnous, Clarias, Saccorbranchus and Ophiocephalus are some of
the actinopterygians, which posses, such organs.

 Lungs develop as a bilobed diverticulum from the floor of the pharynx

posterior to the last gill pouch. They are invaginations of highly vascular
respiratory surface. In higher forms the connection between lungs and
pharynx lengthens and is called the trachea. The upper part of the
trachea becomes modified as a larynx or voice box, the walls of which
are supported by skeletal elements derived from the visceral arches.

 Lungs of lower vertebrate are rather simple, vascular sacs but in higher
vertebrates the walls become subdivided into numerous pocket-like air
spaces. Reptile lungs have vascular partitions known as faveoli. The
divisions become more and more complex in the higher classes of
vertebrates and reach the highest degree of branching in mammals
where the ends of branches form extremely thin walled vasularised blind
ends known as alveoli where the gas exchange actually occurs . The
lungs of birds are complicated in that they are connected to air sacs,
which penetrate among the viscera and even enter the hollow bones.
Birds have a very efficient one-way -flow of air and they can exploit high
altitudes during flight. Instead of alveoli or faveoli they have air tubes
and air capillaries in the lungs where the gas exchange takes place.

 The mechanism employed for inflating and deflating lungs differ in

different vertebrates. The most complex condition is encountered in
mammals, in which the lungs lie in separate pleural cavities partitioned
from the abdominal cavity by muscular diaphragm.

 In vertebrate the main voice producing apparatus are the larynx and
syrinx. Larynx is present in most amphibians, reptiles and mammals but
136 the syrinx is present only in birds.
Unit 4 Respiratory System

4.6 TERMINAL QUESTION

1. Define external and internal respiration.

2. What are the peculiar features of respiratory system of agnathans?

3. Describe the structure of the respiratory system of cartilaginous fishes

and state how does it differs from that of bony fishes.

4. Briefly discuss the mechanism of pulmonary respiration in frog. How is it

different from reptilian mechanism of respiration?

5. How is the respiratory system of birds modified to meet their high

oxygen requirement? How is it different from the respiratory system of
mammals?

4.7 ANSWERS
Self-Assessment Questions
1. (a) F; (b) T; (c) F; (d) F.

2. i) (a) Seven, (b) passive, active; (c) mandibular, hyoid;

(d) Counter current; (e) Interbranchial septum.

ii) (a) False: only elasmobranch gills are supported by interbranchial

septa bony fishes lack it; (b) True; (c) False: they don’t have
external gill slits but an operculum covering a common opening;
(d) True.

3. i) (a) iii; (b) iv; (c) v; (d) i; (e) ii.

ii) (a) disappear; (b) urodeles; (c) do not; (d) disappear.

4. (a) True; (b) False; (c) False; (d) True; (e) True; (f) True;
(g) True; (h) False.

5. a) three, two; (b) nine; (c) active, passive; (d) approximately

500ml; (e) syrinx.

Terminal Questions
1. Refer to Introduction.

2. Refer to Subsection 4.3.2.

3. Refer to Subsection 4.3.3 and compare the two with reference to the gill
structure and ventilation.

4. Refer to Subsections 4.4.1 and 4.4.2.

5. Refer to Subsections 4.4.3 and 4.4.4 and discuss the differences in

terms of respiratory structure and method of ventilation.

137