Ichthyology and Fish Diseases

UNIT II
Structure of Gill & Accessory Respiratory organs
in Fishes
 Contents
 Introduction
 Gills in Fishes
 Spiracles
 Structure of gill
Gross anatomy of gill
Gas exchange over lamellae
 Pseudobranch
Types
 Accessory respiratory organs
Types of accessory
Important modifications in some species
 Swim bladder
 Acknowledgements
 Introduction
– Gills are main respiratory organs in fish. Besides the gills, skin, air bladder and
accessory organs also perform the function of gas exchange in fishes.
– The gills are situated in the pharynx whose lateral walls are perforated by slit like
apertures,, the first of which is situated between mandibular and the hyoid arch
and is called spiracle.
– The expansion and contraction of the pharynx wall and gill silt assist the pumping
of water
– When water is passed over the gills, oxygen is absorbed and carbon dioxide and
ammonium is exhaled.
– 75% of the ammonia excreted by the fish is through the gills.
– The gills also help the fish osmoregulate (equalize body pressures).
– They are covered by a bony flap known as the operculum.
– Elasmobranchs have 5 pair of gill slits/ septal gills.
– Bony Fishes and Chimaeras have four pairs of covered gills.
– Lampeys posses 7 pair of pouched pharyngeal gills
Gills in Fishes

Fig. 1
--

Fig. 2

-- For many bony fishes,

the gill opening is
covered by an
operculum structure
of the gills
 Spiracles
The first pair of gill slit of cartilaginous fish is modified into spiracles (Fig. 3).
It locates on the dorsal surface of ray and skates.
Spiracles are useful for the types of rays that bury themselves in the ocean bottom
because they allow them to breathe without the aid of exposed gills.
Skates, cartilaginous fish that have a flat body and wing-like pectoral fins attached
to their head, and stingrays sometimes use spiracles as their primary method of
 
breathing, bringing oxygenated water into the gill chamber where it is exchanged for
carbon dioxide.

Fig. 3 Spiracles in shark and skate

 Structure of a gill
Gills are formed by outgrowths from the anterior and posterior wall of each gill slit
Healthy gills are bright red and do not stick together.
-Fish gills are supported by cartilaginous or bony structure, the gill arches.
--
 Each gill arch bears two row of slender fleshy projections called gill
filaments.
Gill rakers projects along the inner surface of the gill arch; prevent food
particles from entering and injuring the gill slits, or may be specialized for
filtering feeding. Fig. 4

Fig. 4
  Gross Anatomy
This is the gross anatomy of the gill arches that holds the gill
filaments, that in turn have a number of gill lamellae.
Each gill arch havs a skeletal component that is important for holding
the gill filaments, As you can see in the figure, the blood vessels coming
from the heart (in blue) and going to the body (in red) runs through the
gill arch.
The water flows between the gill arches, through the gill filaments,
passing each gill lamella. This is a effective way of increasing the total
surface area for the exchange.
The total area is the number of lamellae times the surface area of
each lamella. One important aspect is that not all the gill filaments are
exposed to water at rest, which minimizes the problems the fish have
with osmotic differences between the blood (body) and the water.
Each gill filament contains
many rows of thin plates or
dishes called lamellae contains
capillaries; can greatly increase
the respiratory surface.
The number of lamellae is
higher in active swimmers

Fig. 5
 --
Gas exchange over gill lamellae
 The lamellae have a rich blood supply so that a steep concentration
gradient can be maintained between the blood in the lamellae and the
water through.
 Hence, oxygen diffusing into the blood is rapidly removed by the
circulating blood supply and more oxygen is able to difuse into the blood. 
 Another way in which a steep concentration gradient is maintained is by
ensuring water flows in one direction only. 
 The fish opens its mouth to let water in, then closes its mouth and forces
the water through the gills and out through the operculum (gill cover).
 This allows for more efficient gas exchange than if the water had to go in
and out the same way. This is important for fish becaus of the low oxygen
concentration in water. 
 Fig. 5 For gaseous exchange Wter
enters through mouth and flows
over gill lamellae for exit.

Fig. 6: Heart ventral aorta, afferent

brancial artery Gill arch efferent
branchial artery ,dorsal aorta
 Pseudobranch
 The pseudobranch is the vestige of the primary gill arch in teleosts,
positioned in front and toward the back of the operculum (Fig. 7).
 It’s location varies. With some types of fish it is found beneath the skin,
with others it may hang freely.
 Absent in elasmobranchs and cyclostomes, and also in a few teleosts for
example the eel and other related fish. It’s function remains largely
unknown.

Continue..

Fig. 7 location of Pseudobranch

 Pseudobranch
 The term ‘pseudobranch’ was used for a hemibranch which has lost its respiratory
function.
 It may be free or covered with a layer of mucous membrane and consists of a
series of gill filaments. It develops early in the embryo and may be respiratory in
embryonic stage but not in the adult.
 The pseudobranch in teleosts receives oxygenated blood directly from the dorsal
aorta or an efferent branchial artery and has a vascular connection with the
internal carotid artery.
 It is present in a large number of teleosts and is free or glandular; but is absent in
cat fishes (as the Wallago, Mystus), the eels and species belonging to Gymnarchus
and Cobitis.
 A teleostean pseudobranch is reported to represent the posterior mandibular
hemibranch.
 In sharks, rays, chondrosteans and holosteans, the pseudobranch may be derived
from the posterior mandibular hemibranch, but in teleosts it is derived from the
posterior hyoidean hemibranch.
 Types of Pseudobranch:
• Pseudobranch varies in shape, size and location in various teleosts, but its blood supply and
innervation suggest a common plan. It is either attached to the operculum, as in Hilsa ilisha or
is deeply embedded in the connective tissue of the roof of the buccal cavity, as in Channa.
• According to Bertin (1958), pseudobranch may be of the following types:
• (1) Free Type:
• Free pseudobranch with a row of filaments and secondary lamellae is seen in Clupeidae (Hilsa,
Gudusia), Syngnathidae, Labridae, Pleuronectidae and Lophidae. A free pseudobranch (as in
Hilsa, Gudusia) is exposed to water and retains the basic gill structure, having filaments,
lamellae and pillar cells.
• (2) Covered Type:
• In several species as the Glossogobius, Gadus, Cyprinus, Catla, Phoxinus, etc. it is a gill-like
structure but is covered with the opercular membrane and connective tissue.
• (3) Glandular Type:
• The pseudobranch is deeply embedded in the thick connective tissue of the opercular cavity or
the buccopharynx as in Channa, Anabas and Notopterus.
• The covered pseudobranch loses its contact with water, and its filaments and lamellae are
fused, which in extreme cases form a glandular structure.
 Accessory Respiratory organs
 Sometimes the fishes of the tropical freshwaters and hill-streams
develop accessory respiratory organs to meet extra demand for
oxygen.
 Accessory respiratory organs enable the fishes to live in oxygen-
deficient water, to aestivate over prolonged droughts in dry summer,
to take excursions on land or simply to meet extra demand for oxygen.
 To overcome these adverse situations, accessory respiratory organs
functionable in aquatic and/or aerial environment have been
developed in fishes.
 So the development of such structures is essentially adaptive in native.
Some accessory organs sub serve aquatic respiration, while others
aerial respiration (Fig. 17.8).
Accessory Respiratory Organs in Teleost Fishes
 Types of Accessory Respiratory Organs
• Several types of accessory respiratory organs have been evolved in different
species of fishes. 
• These accessory respiratory organs of fishes are as follows:
• 1. Skin or Integument:
• In the eel, Anguilla anguilla, Amphipnous cuchia and in Periophthalmus and
Boleophthalmus, the skin is highly vascular and serves for exchange of gases as in frog,
when the fish is out of water. These fishes habitually leave the water and migrate from one
place to another through damp vegetation. During this period, the moist skin serves as an
important organ in respiration. They can respire cutaneously both in air and in water.
• Since Amphipnous and Mastacembelus live in oxygen deficient stagnant water, the skin is of
little use for respiration but it plays an important role in extracting oxygen from air, when
the fishes are exposed in drying up muddy ponds, or when fish is moving out of water. The
glandular secretions of the skin protect it from desiccation in the air.
• Median fin folds of many fishes are supplied with numerous blood vessels that help in
cutaneous respiration. Besides, the highly vascular opercular folds of Sturgeons and many
cat fishes serve as accessory respiratory structures.
2. Bucco-Pharyngeal Epithelium:

 In most of the fishes, the epithelial lining of buccal cavity and pharynx is usually
highly vascular and permeable to gases in water.
 It may remain simple or may develop folds, pleats or tongues projecting into the
buccal cavity and pharynx to make it an efficient respiratory organ.
But in mudskippers (Periophthalmus and Boleophthalmus) the highly vascularised
buccopharyngeal epithelium helps in absorbing oxygen directly from the
atmosphere. These tropical fishes leave water and spend most of the time skipping
or walking about through dampy areas particularly round the roots of the
mangroove trees. The old idea that the mudskippers use the vascular tail as the
respiratory organ is not supported by recent ichthyologists.
3. Gut Epithelium:
 In several fishes epithelial lining of certain parts of alimentary canal becomes
vascular and modified to serve as a respiratory organ. It may be just behind
stomach (Misgurus fossilis) or intestine (Lepidocephalus guntea, Gobitus (giant
loach of Europe) or rectum (Callichthyes, Hypostomus and Doras).
 Fresh air is drawn through mouth or anus and after gaseous exchange the gas is
voided through the anus. In these fishes the wall of the gut is modified to perform
the respiratory function. The walls of the gut in these areas become thin due to
the reduction of muscular layers. . 
4. Outgrowths of Pelvic Fins:
 In American lung fish, Lepidosiren, during breeding time, the pelvic fins of male
become enlarged and grow filamentous vascular outgrowths which provide fresh
oxygen to the guarded eggs.
• epithelium) is present on each side of the roof of the pharynx.

5. Opercular Chamber Modified for Aerial Respiration:

 In some species, the inhaled air is passed through the gill-slits into the opercular chamber
where it is stored for some time. The opercular chamber becomes bulged out in the form
of two little balloons in the hinder region of the head and after sometimes its walls
collapse and the air is passed out through the small external branchial opening. The
membrane lining the opercular chamber becomes thin and highly vascular to allow
exchange of gases. This is seen in Periophthalmus and Boleophthalmus.

6. Branchial Diverticula:
 The outgrowths from gill-chambers form more complicated aerial accessory respiratory
organs than the simpler pharyngeal outgrowths in other fishes. Such air breathing organs
are present in Heteropneustes, Clarias, Anabas, Trichogaster, Macropodus, Betta, etc.
7. Pharyngeal Diverticula:
 Pharyngeal diverticula are a pair of simple sac-like outgrowths of pharynx, lined by
thickened vascular epithelium and extending above the gills. In Channa
( Ophiocephalus), the accessory respiratory organs are relatively simpler and
consist of a pair of air-chambers (Fig. 17.8).
 These are developed from the pharynx and not from the branchial chambers as
seen in others. The air-chambers are lined by thickened epithelium which is highly
vascularised. The air-chambers are simple sac-like structures and do not contain
any structure. These chambers function as the lung-like reservoirs. In Channa
striatus, the vascular epithelium lining the chambers becomes folded to form some
alveoli. The gill-filaments are greatly reduced in size.
 In cuchia eel, Amphipnous cuchia, the accessory respiratory organs consist of a
pair of vascular sac-like diverticula from the pharynx above the gills (Fig. 17.8).
These diverticula open anteriorly into first gill-slit.
 These diverticula function physiologically as the lungs. The gills are greatly reduced
and a few rudimentary gill-filaments are present on the second of the three
remaining gill-arches. The third gill-arch is found to bear fleshy vascular
(respiratory) epithelium.
 In Periophthalmus also, a small, shallow pharyngeal diverticulum lined with
respiratory epithelium (vascular
 Important Modifications in Some Species
(a) Heteropneustes Fossilis (= Saccobranchus): This Indian catfish has a pair of long,
tubular and dorsally situated air-sacs, arising posteriorly from gill-chambers and
extending almost up to the tail. They are highly vascular. The air is drawn in and
expelled out through pharynx.
(b) Anabas Testudineus: The Indian climbing perch has two, spacious, suprabranchial
cavities as dorsal outgrowths of the two gill-chambers. Each cavity contains a special
labyrinthine organ formed of much folded, concentric bony plates developed from
the first epibranchial bone and covered with thin vascular mucous membrane.
Margins of these plates are wavy and the plates are covered with vascular gill-like
epithelium.
Each branchial outgrowth communicates freely not only with the opercular cavity but
also with buccopharyngeal cavity. Air is drawn through mouth into suprabranchial
cavities and expelled through opercular opening. The fish is so dependent on
atmospheric oxygen that it will drown if denied access to surface to gulp air.
(c) Trichogaster Fasciatus: The accessory respiratory organs in this species consist of a
suprabranchial chamber, a labyrinthine organ and the respiratory membrane. The
suprabranchial chamber is situated above the gills on either side as in Anabas,
communicates with the pharynx by means of inhalent aperture and with the
exterior through the opercular chamber by means of an exhalent aperture.
 The labyrinthine organ develops from the epibranchial of the first gill-arch and is
simpler in structure than that of Anabas. It is in the form of a spiral organ
possessing two leaf-like expansions and is composed of loose connective tissue
covered by a vascular epithelium.
 The respiratory membrane lining the air-chamber and covering the labyrinthine
organs consist of vascular and non-vascular areas, of which the former possesses a
large number of ‘islets’ containing parallel blood capillaries. The islets are believed
to be derived from the secondary lamellae of a typical gill-filament.
 Clarias Batrachus:
 The Indian cat fish, Clarias batrachus has the most complicated accessory respiratory
organs.
 The accessory air-breathing organs of this fish consist of:
 (i) The suprabranchial cavity or chamber,
 (ii) The two beautiful ‘rosettes’ or air-trees or arborescent organs or dendritic organs,
 (iii) The ‘fans’ and
 (iv) The respiratory membrane.
 The suprabranchial chamber lies above the gills and is divided into two cup-like
compartments and is lined by a highly vascular respiratory membrane.
 Two beautiful ‘rosettes’ or dendritic organs are present on each side and are supported by
epibranchials of the second and the fourth branchial arches. The first of these is smaller in
size and lies in the anterior compartment. Each is a highly branched tree-like structure
supported by cartilaginous internal skeleton. The terminal knobs or bulbs of each dendritic
organ consist of a core of cartilage covered by vascular epithelium showing eight folds in it.
 Swim Bladder/Air-Bladder:
 Swim-bladder of higher bony fishes (teleosts) is essentially a hydrostatic organ. But in
lower bony fishes (dipnoans and ganoids), the air-bladder acts like a lung to breathe
air and is truly an accessory respiratory organ. The wall of bladder is vascular and
sacculated with alveoli. In Amia and Lepidosteus, the wall of the swim-bladder is
sacculated and resembles lung.
 In Polypterus, the swim-bladder is more lung-like and gets a pair of pulmonary
arteries arising from the last pair of epibranchial arteries. The swim-bladder in
dipnoans resembles strikingly the tetrapod lung in structure as well as in function. In
Neoceratodus, it is single but in Protopterus and Lepidosiren it is bilobed (Fig 6.87).
 The inner surface of the ‘lung’ is increased by spongy alveolar structures. In these
fishes, the lung is mainly respiratory in function during aestivation because the gills
become useless during this period.
 Like that of Polypterus, the ‘lung’ in dipnoans gets the pulmonary arteries from the
last epibranchial arteries. The swim-bladder of feather tail, Notopterus notopterus
has a wide pneumatic duct and a network of blood capillaries covered by a thin
epithelium in its wall. This helps in exchange of gases.
http://www.biologydiscussion.com/fisheries/fish/swim-bladder-development-structure-a
nd-types-fishes/40812
 Functions of Accessory Respiratory Organs:
 The fishes possessing such respiratory organs are capable of living in
water where oxygen concentration is very low. Under this condition
these fishes come to the surface of water to gulp in air for
transmission to the accessory respiratory organs.
 If these fishes are prevented from coming to the surface, they will die
due to asphyxiation for want of oxygen. So the acquisition of accessory
respiratory organs in fishes is an adaptive feature.
 Further it has been observed that the rate of absorption of oxygen in
such organs is much higher than the rate of elimination of carbon
dioxide.
 Hence, it is natural that the gills excrete most of the carbon dioxide.
Absorption of oxygen appears to be the primary function of the
accessory respiratory organs.

THE END
 This PPT is prepared from available free
access internet material, Figures and Text.
The sources/authors are duly cited and
highly acknowledged.

THANKS!!!