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Respiration is the process of obtaining oxygen from the external environment & eliminating
1. External respiration - oxygen and carbon dioxide exchanged between the external
environment & the body cells
2. Internal respiration - cells use oxygen for ATP production (& produce carbon
dioxide in the process)

Adaptations for external respiration:

1 - Primary organs in adult vertebrates are external & internal gills, swim bladders or
lungs, skin, & the buccopharyngeal mucosa
2 - Less common respiratory devices include filamentous outgrowths of the posterior
trunk & thigh (African hairy frog), lining of the cloaca, & lining of esophagus

Respiratory organs:
Cutaneous respiration
a. respiration through the skin can take place in air, water, or both
b. most important among amphibians (especially the family Plethodontidae)


Cartilaginous fishes:
1. 5 ‘naked’ gill slits
2. Anterior & posterior walls of the 1st 4 gill chambers
have a gill surface (demibranch). Posterior wall of last
(5th) chamber has no demibranch.
3. Interbranchial septum lies between 2 demibranchs of a
gill arch
4. Gill rakers protrude from gill cartilage & ‘guard’
entrance into gill chamber
5. 2 demibranchs + septum & associated cartilage, blood
vessels, muscles, & nerves = holobranch

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

Bony fishes (teleosts):
1. usually have 5 gill slits
2. operculum projects backward over gill chambers
3. interbranchial septa are very short or absent

1. 6 - 15 pairs of gill pouches
2. pouches connected to pharynx by afferent branchial (or gill)
ducts & to exterior by efferent branchial (or gill) ducts

Larval gills:
1. External gills
a. outgrowths from the external surface of 1 or more gill
b. found in lungfish & amphibians
2. Filamentous extensions of internal gills
a. project through gill slits
b. occur in early stages of development of elasmobranchs
3. Internal gills - hidden behind larval operculum of late anuran tadpoles

Swim bladder & origin of lungs - most vertebrates develop an outpocketing

of pharynx or esophagus that becomes one or a pair of sacs (swim bladders or
lungs) filled with gases derived directly or indirectly from the atmosphere.
Similarities between swim bladders & lungs indicate they are the same organs.
Vertebrates without swim bladders or lungs include cyclostomes, cartilaginous fish, and a few
teleosts (e.g., flounders and other bottom-dwellers).

Swim bladders:
1. May be paired or unpaired (see diagram above)

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

2. have, during development, a pneumatic duct that
usually connects to the esophagus. The duct remains
open (physostomous) in bowfins and lungfish, but
closes off (physoclistous) in most teleosts.
3. Serve primarily as a hydrostatic organ (regulating a
fish's specific gravity)
4. Gain gas by way of a 'red body' (or red gland); gas is
resorbed via the oval body on posterior part of
bladder may also play important roles in:
a. Hearing - some freshwater teleosts (e.g., catfish, goldfish, & carp) 'hear' by way
of pressure waves transmitted via the swim bladder and small bones called
weberian ossicles (see diagram below)
b. sound production - muscles attached to the swim bladder contract to move air
between 'sub-chambers' of the bladder. The resulting vibration creates sound in
fish such as croakers, grunters, & midshipman fish.
c. Respiration - the swim bladder of lungfish has number subdivisions or septa (to
increase surface area) & oxygen and carbon dioxide is exchanged between the
bladder & the blood

Lungs & associated structures

1. Larynx
a. Tetrapods besides mammals - 2 pair of cartilages: artytenoid & cricoid
b. Mammals - paired arytenoids + cricoid + thyroid + several other small cartilages
including the epiglottis (closes glottis when swallowing)
c. Amphibians, some lizards, & most mammals - also have vocal cords stretched
across the larynx.
2. Trachea & syrinx
a. Trachea
i. usually about as long as a vertebrates neck (except in a few birds such as
ii. reinforced by cartilaginous rings (or c-rings)
iii. splits into 2 primary bronchi &, in birds only, forms the syrinx at that

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

3. Lungs
a. Amphibian lungs
i. 2 simple sacs
ii. internal lining may be smooth or have simple
sacculations or pockets
iii. air exchanged via positive-pressure ventilation

b. Reptilian lungs
i. simple sacs in Sphenodon & snakes
ii. Lizards, crocodilians, & turtles lining is septets,
with lots of chambers & sub chambers
iii. air exchanged via positive-pressure ventilation
a. Avian lungs - modified from those of reptiles:
i. air sacs (diverticula of lungs) extensively distributed throughout most of
the body
ii. arrangement of air ducts in
lungs ----> no passageway is a
iii. air flow through lungs
(parabronchi) is unidirectional
a. Mammalian lungs:
i. multi chambered & usually
divided into lobes
ii. air flow is bidirectional:
6. Trachea <---> primary bronchi <--->
secondary bronchi <---> tertiary bronchi <---
> bronchioles <---> alveoli

7 . Air exchanged via negative pressure ventilation, with pressures changing due to
contraction & relaxation of diaphragm & intercostals muscles

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016


Gills in Protochordates

A large and sieve-like pharynx in majority of these animals performs dual function of
respiration and trapping food particles which are brought in through the current of water. The
primitive pterobranch hemichordates (Cephalodiscus and Rhabdopleura) have either no gill slits
or have very few and sport tentaculated arms, which other than food gathering, also function as
efficient respiratory organs. Balanoglossus possesses a large pharynx having as many as 700
pairs of gill slits, which appears to be a necessity in the burrowing habitat of the animal.
The free-living urochordates, such as Salpa and Doliolum do not possess many stigmata
or gill slits as their entire body is permeable to oxygen but in the sedentary ascidians pharynx is
prominently enlarged and perforated with no less than 200,000 stigmata for filter-feeding.
Cephalochordates use pharynx for both filter-feeding and respiration and hence carry
150-200 pairs of gill slits.

Respiratory organs of Cyclostomes

Agnathans have 6-15 pairs of gill pouches, which are lateral extensions of pharynx and
contain gill lamellae within. Cyclostomes are called marsipobranchs, which means “pouched
gills”, since the gill lamellae are housed in gill pouches. The hagfish, Myxine has only 6 pairs of
gill pouches whose ducts join together and open to the exterior by a single pair of openings,
while Bdellostoma carries 6-15 pairs of gill pouches that vary in different species and open to the
outside independently. In Myxine behind the gill pouches there is a single pharyngocutaneous
duct on the left side, which is a modified gill pouch which drains excess water that fails to enter
the gill pouches. The lamprey, Petromyzon, has 8 embryonic and 7 adult paired gill pouches that
open to the exterior by independent openings.

Respiratory organs in Elasmobranchs

Most Elasmobranchs possess 5 pairs of gill slits and a pair of spiracles. There is no
operculum covering the gill slits in cartilaginous fishes. A demibranch is a bunch of gill
lamellae attached on one side of the interbranchial septum. Hence, there are altogether 9 pairs of

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

demibranchs in Elasmobranchs. Between the two demibranchs lies the interbranchial septum,
which is supported by gill cartilages. Anterior to the first gill slit is a spiracle or pseudobranch. In
free swimming sharks and dogfishes water generally enters through the mouth.
Blood to the gills is supplied by five pairs of afferent branchial arteries coming from
ventral aorta and hence they bring deoxygenated blood from heart. Blood is then oxygenated in
gills and is collected by the loops of four pairs of efferent branchial arteries and carried to the
paired dorsal aorta, the two sides of which meet posteriorly to form single median dorsal aorta
that supplies oxygen-rich blood to the whole body.

Gills of bony fishes

In bony fishes gills are covered with an operculum that is made of flattened skeletal
plates and there is no spiracle as in Elasmobranchs. There are 4 pairs of gill pouches, each
containing two demibranchs, making the total number of demibranchs in bony fishes as 8 pairs
or four pairs of complete gills or holobranchs. Teleosts always breathe with their mouth open and
eject expiratory water by opening operculum. Gills in Chondrostei, Holostei and the lungfish
Neoceratodus exhibit partial reduction in their interbranchial septa, which happens to be
somewhat intermediate condition between Elasmobranchs and teleosts.


External gills develop from the outer wall of pharynx or from the exposed portion of
branchial arch. They occur in larval lampreys, few larval fishes, Polypterus, lungfishes, some
larval teleosts and all larvae and some adults of amphibians. There is a single pair of larval gill in
the chondrosteian bony fish, Polypterus, which has a long axis carrying gill lamellae. The
African and South American lung fishes possess 4 pairs of feathery external gills. The larval
forms of some amphibians and some adult urodeles possess external gills which arise simply as
folds of skin on the surface of the III, IV and V branchial arches but weakly supported by the
skeletal system. Perennibranch amphibians as Necturus and Proteus retain external gills
throughout life along with 2 or 3 pairs of gill slits, which are functionless as the water does not
pass through pharynx. Instead, gills are waved in water by means of muscles attached at the base
of gill axis for respiration. The larvae of limbless amphibian, Caecilia, have a pair of
exceptionally large leaf-like gills with profuse blood supply. Salamanders that inhabit hill
streams, e.g. Eurycea and Salamandrina, which belong to family Plethodontidae have neither
gills nor lungs for respiration and survive only on cutaneous respiration.


Barring agnathans, cartilaginous fishes and few bottom dwelling teleosts, all fishes carry
a gas-filled air bladder on the dorsal side of the gut, which serves as hydrostatic organ. On the
ventral side of the bladder there occurs a highly vascularised area called red gland that is
supplied by intestinal artery and portal vein and which has unique capability of extracting free
oxygen from the blood and release it into the air bladder in order to make it inflate. Small pouch-
like diverticula called oval that can be closed or opened by sphincter muscles is the site of
reabsorption of gases. Secretion and absorption of gases in swim bladder occurs under the
control of autonomic nervous system, based on the depth at which a fish is swimming.

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

In Cypriniformes (Teleosti), a series of four small bones (tripus, intercalarium,
scaphium and claustrum), derived from the first three vertebrae and called Weberian Ossicles,
connect the anterior end of air bladder with the sinus impar of membranous labyrinth. Sound
vibrations received by air bladder from the surrounding water are conveyed to the internal ear
through this unique apparatus to bestow some hearing ability to these fishes.
In some fishes as for example ganoids, carps and catfishes, a pneumatic duct connects
the air bladder with oesophagus. Such condition is called physostomous (Gr. physo=bag;
stoma=opening). Fishes which do not have such a pneumatic duct connecting the air bladder are
called physoclistous (Gr. physo=bag; clista=closed).
The comparative study of air bladders in different groups of fishes and striking similarity
between the swim bladder and lung suggest a phylogenetic relationship between the two. The
conventional belief is that lungs evolved from the air bladder of fishes. However, recent
evidences point to the contrary that lungs evolved first in fishes for supplementing oxygen from
air and then they got transformed into swim bladder as the oxygen concentration in water

Accessory Respiratory Organs in Fishes

Many species of fishes developed breathing organs other than gills for supplementing
deficiency of oxygen in water. These are as follows:

Dendritic Organs

They are also called arborescent organs as they are highly vascularised tree-like,
branched structures produced by the second and fourth gill arches and located in the
suprabranchial chamber, posterior to the gills. Paired gill fans at the opening of branchial
chamber force air over the dendritic organs as the fishes gulp air. Dendritic organs are found in
catfishes such as Clarias.

Labyrinthine Organs

These are rosette-like concentric plates of tissue present in the suprabranchial chamber
of climbing perch (Anabas), Trichogaster, Osphromanus and Polycanthus. Respiration takes
place when these fishes gulp air. Perches can migrate from one pond to another by breathing air
through labyrinthine organs and using pectoral fin spines to walk on land.

Pneumatic Sac

It is a tube like extrabranchial diverticulum that extends up to tail in some cat fishes such
as Heteropneustes, which can survive out of water for considerable time using these organs for
air breathing.

Air Chamber

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

Air chamber is a small, highly vascularised sac located behind the gills of some fishes,
e.g. Ophiocephalus, Macropodus and cuchia eel (Amphipnous). These fishes can gulp air and use
it as air breathing organ.

Buccopharyngeal epithelium

Mud skippers (Periophthalmus, Balaeophthalmus) possess vascularised buccopharyngeal

epithelium and also a respiratory tail. They skip around in swamy areas, breathing air by
buccopharyngeal epithelium or keep their tail in water for aquatic respiration.


Eels (Anguilla) breathe through skin while migrating from the American and European
rivers to Sargasso Sea in Bermuda. As much as 60% exchange of gases takes place through the
highly vascularised skin.

Gut epithelium

Fishes such as Callichthys, Hypostomus, Doras, Misgurnus, Cobitis can suck and release
water through anus and exchange of gases can take place in the rectal lining. In giant loach
(Cobitis) and Misgurus lining of stomach and intestine is used as respiratory organ.

Lungs of Polypterus and the ganoid fish Calamoichthys are asymmetrical and connected
by pneumatic duct on the ventral side of pharynx. The blood is supplied to lung by pulmonary
artery that emerges off the 6th aortic arch, but unlike in lungfishes venous blood returns to
hepatic vein.
Lungs of Dipnoi (Choanichthys) are bilobed or paired as in Protopterus (African lung
fish) and Lepidosiren (South American Lung fish) and are connected to oesophagus via a
pneumatic duct. But the Australian lung fish (Neoceraodus) has a single lung that is used as
hydrostatic organ.
In tetrapods, embryonic lungs arise from pharyngeal wall as a hollow mid-ventral
evagination that subsequently bifurcates to form two lungs that carry an envelope of peritoneum.


Lungs of amphibians are two simple sacs, narrow and elongated in urodeles and bulbous
in anurans enclosed in a single peritoneal membrane and supplied by pulmonary arteries and
drained by pulmonary veins. Left lung in limbless amphibians is rudimentary. Lungs are
vestigial in salamanders inhabiting hill streams where in fast flowing water buoyancy would not
be a desirable trait.
Amphibians lack ribs and hence use floor of the buccal cavity to force air in and out of
the lungs. Frogs and toads modify 2nd, 3rd and 4th visceral arches to produce a plate-like
hyobranchial apparatus that lies in the floor of oral cavity and is connected to squamosal bone of
skull by petrohyal muscle and to sternum by sternohyal muscle. One breathing cycle is
completed in four steps in anurans that is affected by contraction of these two muscles.

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

Breathing in frog requires much faster movement of hyoid plate as compared to lungs and
considerable amount of gas exchange takes place in bucco-pharyngeal region too. Cutaneous
respiration also contributes to major part of oxygen supply to the body of amphibians.


Lungs are narrow and elongated in snakes and lizards extending up to two-third of the
body cavity but are more bulbous in turtles and crocodiles. The left lung is rudimentary in
limbless lizards and snakes. There are well formed alveoli in lungs which are housed securely in
a pair of pleuro-peritoneal cavities. Breathing in snakes and lizards is carried out by a
combination of hyoid plate, nostril valves and ribs or only by the movement of rib cage, while
tortoises and turtles make use of muscles surrounding peritoneal membranes. Crocodiles are the
only living reptiles that possess a muscular diaphragm for breathing as do the mammals.


Lungs are secured into pleural cavities and extend into membranous air sacs that occupy
all available space in the body cavity and also penetrate into bone marrow cavities. This makes
the bones pneumatic in birds and help to reduce body weight which is so necessary in flight.
Majority of birds have 5 pairs of air sacs, namely, cervical at the base of neck; interclavicular
often united across midline; anterior thoracic placed lateral to the heart; posterior thoracic
within the oblique septum and abdominal within the abdominal cavity. Sometimes there are also
axillary air sacs near the pectoral muscles. The flight muscles inflate and deflate the air sacs like
bellows with each stroke of wings.

Air duct system is unique in birds as there are no alveoli as found in reptiles and
mammals. Trachea divides into two bronchi which enter the lungs and branch into mesobronchi
that again divide to form parabronchi. From each parabronchus, bunches of air capillaries arise
which loop back into their own lumen to form anastomosis that eventually leads into the air sacs.
Air capillaries are minute and only one cellular layer thick and contain respiratory epithelium
and rich network of blood capillaries.
During inspiration and expiration, air passses through the air capillary anastomosis into
the air sacs and back twice making it a double respiration.


Mammalian lungs increase efficiency by increasing the surface area of respiratory

epithelium of alveoli whose number goes up to millions and lungs become almost like semisolid
sponge with little empty lumen inside. Lungs are enclosed in double peritoneal membranes, the
outer parietal pleura and inner visceral pleura that enclose the fluid-filled pleuro-peritoneal
cavity in between.
Trachea which is commonly known as windpipe opens in pharynx by a slit-like glottis
and posteriorly divides into two primary bronchi that enter lungs and branch off to secondary and
tertiary bronchi which ultimately lead to fine capillaries called bronchioles. Each bronchiole is
connected to several alveoli by alveolar ducts. The number of alveoli in human lungs is

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

estimated to be about 750 million which collectively carry an enormous surface area of about
100 m2 and if stretched.
A muscular diaphragm located between the thoracic and abdominal cavities moves in the
antero-posterior direction and forces air in and out of the lungs. External costal muscles and
internal costal muscles are attached between the ribs and sternum, the former increases the
thoracic space while the latter decreases it to carry out what is known as thoracic respiration or
common panting after strenuous physical exercise.
Whales possess enormous nasal chambers in the head that can store large quantity of air
when diving deep in the sea. Passage of air from nasal chambers to lungs is controlled by a pair
of valves.


Sound producing organs, larynx and syrinx, are associated
with trachea through which air can be forced from lungs into the sound
box to produce sound. Larynx in urodeles is so simple that it has only a
pair of lateral cartilages, called Guardian cartilages that surround the
glottis and this apparatus is incapable of producing any sound in these
Larynx of frog is made of a cricoid cartilage which is a
modification of the first tracheal ring and a pair of arytenoid cartilages,
which support a pair of vocal cord that vibrates to produce sound.
Males of frogs and toads in addition possess a pair of vocal sacs which
are evagination of oral cavity and serve as resonance chambers to
amplify sound.
Reptiles are silent animals but possess larynx, albeit without a vocal cord, in the absence
of which they can at best produce a hissing sound.
Birds inherited a rudimentary larynx from their reptilian ancestors and hence evolved a
secondary sound producing organ called syrinx located at the junction of trachea and bronchi
and hence called bronchotracheal type. The tympanic chamber has a bony ridge at base called
pessulus that supports the unpaired membrana semilunaris, which vibrates to produce sound.
In mammals larynx consists of paired arytenoid cartilages that support vocal cord. Base
of the sound box is made of a ring-like cricoid cartilage and the unpaired thyroid cartilage forms
the surrounding walls of the tympanic chamber. A pair of fleshy vocal cords is stretched across
the cephalic part of the vocal chamber supported by the paired cartilages of Santorini. The cord
vibrates to produce sound that is modulated in the oral cavity.

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016
Dr.C.V.Narasimha murthy . Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016