Fish respiration

HOW DO FISH RESPIRE?

Adult fish have a pair of gills. Each gill is covered by a boney lid (removed from the
picture). A fish draws in water by closing the lid over its gills and opening its mouth.
When the fish closes its mouth and opens the gill lid the water is forced out and over the
respiratory surfaces of the gill filaments.

HOW IS OXYGEN AND CARBON DIOXIDE EXCHANGED IN THE

GILLS?

In order to live, fish must extract oxygen from the water and transfer it to their
bloodstream. This is done by gills which are richly supplied with blood vessels in order to
act as a respiratory organ. Extracting oxygen from water is more difficult and requires a
greater expenditure of energy than does extracting oxygen from air. Water is a thousand
times denser than air, and at 20 deg C it has 50 times more viscosity than air and contains
only 3% as much oxygen as an equal volume of air. Fishes, therefore, have necessarily
evolved very efficient systems for extracting oxygen from water; some fishes are able to
extract as much as 80% of the oxygen contained in the water passing over the gills,
whereas humans can extract only about 25% of the oxygen from the air taken into the
lungs. The oxygenated water flows through the tinny gill filaments and it exchanges the
carried oxygen for carbon dioxide through a process called diffusion.
WHAT IS THE STRUCTURE OF A GILL?

A large surface area for gaseous exchange means that more oxygen can enter the
bloodstream over a given period of time. A single gill of a bony fish consists of a curved gill
arch bearing a V-shaped double row of gill filaments. Each filament has many minute
folds in its surface, giving it a sort of fuzzy appearance and increasing the amount of
surface area along a given length of filament. Consequently, the surface area of the gills is
commonly 10 to 60 times more than that of the whole body surface.

HOW ARE GILLS EFFICIENT?

1) A short diffusion, or travel, distance for the oxygen increases the rate of oxygen entry
into the blood. The blood traveling in the folds of the filaments is very close to the oxygen-
containing water, being separated from it by a very thin membrane usually 1 to 3 microns
(4/100,000 to 1/10,000 in) or less thick.

2) By using countercurrent circulation in the gill, the blood in the filament folds travels
forward, in the opposite direction to the water flow, so that a constant imbalance is
maintained between the lower amount of oxygen in the blood and the higher amount in the
water, ensuring passage of oxygen to the blood. If the blood were to flow in the same
direction as the water, oxygenated blood at the rear of the gills would be traveling with
deoxygenated water and not only could not extract oxygen from the water but would even
lose oxygen to it.

3) Gills have little physiological dead space. The folds of the filament are close enough
together so that most of the water passing between them is involved in the gas-exchange
process.
 4) Water flows continuously in only one direction over the gills, as contrasted with the
interrupted, two-way flow of air in and out of lungs of mammals.
IN CONCLUSION...
The surface area of the gill filaments is a factor that means death or life to a fish since the
water contains much less oxygen than air therefore fish must have an organ with large
surface area in order to absorb enough oxygen from the water to survive.
This chapter provides a general overview of fish air breathing, vertebrate evolution, and comparative
physiology. Air breathing has persisted throughout the evolutionary history of the fishes and has played
a fundamental role in this group's evolution. Modern air-breathing fishes occur in a variety of freshwater
and marine habitats. Marine air breathers occur on tropical coral reefs, in rocky intertidal zones, and in
marshes and bays. A variety of factors, from the adoption of an amphibious behavior to exploit
resources at the airwater interface, to the periodic aerial exposure imposed by low tides, and even
aquatic hypoxia in certain habitats have all played a role in selection for air breathing among marine
species. This chapter reviews physical principles governing the suitability of air and water as respiratory
media, and examines the habitats of extant air-breathing fishes. The chapter also defines terms and
concepts related to aerial and aquatic gas exchange and bimodal respiration. The chapter provides an
operational definition of fish air breathing along with a discussion of the types of observations,
experimental evidence, and background knowledge required to evaluate the presence of air breathing in
a species. The chapter also presents a general classification of the types of air breathing fishes.

This chapter focuses on the structure of fish air-breathing organs (ABO). The principal requirements for
an ABO are the acquisition, retention, and absorption of aerial oxygen. Few structures other than the
mouth and jaws can be used to capture air. In terms of holding inspired air, spacious cavities or
compliant spaces with a large blood supply and potentially modifiable vascular surfaces either already
occur or can be developed in the mouth, branchial region, or digestive track. Thus, the form taken by
the ABOs of modern fishes is characterized by either evolutionary parallelism or convergence driven by
limitations of structure, space, and surface area. This chapter discusses in detail the comparative
morphometrics of ABOs, their air-ventilation mechanics, tidal volumes, and the timing and sequence of
air inhalation and expiration. The chapter discusses the gill and skin morphology in relation to air
breathing. The chapter also presents a simplified ABO classification scheme that groups all the primitive
fishes possessing lungs or respiratory gas bladders.

This chapter examines the influence of aerial gas exchange on the blood-respiratory properties of air
breathing fishes. With few exceptions, fishes have a tetrameric Hb contained within a nucleated
erythrocyte. To a large extent, both the Hb concentration and Hb-O2 affinity of air-breathing fish blood
are functionally linked to circulatory specializations for conserving aerially-obtained O2 and preventing
mixing between the air breathing organ (ABO) efferent and systemic bloods. An elevated blood O2
capacity compensates for the forced unsaturation of O2-rich efferent ABO blood when it mixes with O2-
poor systemic blood in the venous circulation. A high [Hb] can serve both an O2 storage and a blood-
buffering function in air-breathing fishes. Although the pattern of reduced Hb-O2 affinity is seen among
closely related species of non-air-breathing and air-breathing osteoglossids and erythrinids, comparisons
among a diversity of air-breathing and non-air-breathing species fail to verify this as a universal
evolutionary pattern. Research indicates that the degree of branchial isolation enjoyed by a species, that
is, its degree of independence from gill respiration, is a major determinant of Hb-O2 affinity during air
breathing.

Aerial and Aquatic Gas Exchange

This chapter reviews the diverse information that is found under the general subject of air-breathing
fish respiration. Studies of air-breathing fish respiration have identified factors regulating the duration
of the air-breath cycle, determined respiratory capacity, and elucidated respiratory organ function
during bimodal respiration. For most aquatic air breathers, aerial respiration is utilized to the extent
needed to supplement aquatic respiration and sustain normal VO2. For amphibious air breathers, the
complete transition from aquatic to aerial gas exchange ensures a period of total respiratory
independence from water. Air-breath durations differ among species. These are highly variable, and are
affected by several physical factors including aquatic oxygen level and temperature. In situations of
chronic hypoxia, fish may ascend for air as frequently as 1200 times in 24 hours. Light level also affects
air-breath duration as can the presence of water-borne toxins. An effect of aquatic CO2 has also been
suggested for some species. This chapter begins with a historical overview, followed by a comparison of
air breathing cycles and a review of whole-organism respiratory gas exchange. The chapter also
examines the specialized aspects of respiratory organ function. The chapter emphasizes the comparative
physiological aspects of air breathing.

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Gas Bladder in Fishes (With Diagram)

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In this article we will discuss about:- 1. Gas Bladder as a Respiratory Device 2. Blood
Supply of Gas Bladder 3. Histology 4. Gas Bladder in Sound Production 5. Gas Bladder in
Sound Reception 6. Gas Bladder as a Hydrostatic Organs 7. Filling and Emptying of the
Gas Bladder 8. Secretion of Gas from Blood to Lumen of Bladder 9. Reabsorption of Gas
from Bladder.

Contents:

1. Gas Bladder as a Respiratory Device

2. Blood Supply of Gas Bladder
3. Histology of Gas Bladder
4. Gas Bladder in Sound Production
5. Gas Bladder in Sound Reception
 6. Gas Bladder as a Hydrostatic Organs
7. Filling and Emptying of the Gas Bladder
8. Secretion of Gas from Blood to Lumen of Bladder
9. Reabsorption of Gas from Bladder

1. Gas Bladder as a Respiratory Device:

Gas bladder is one of the characteristic feature of the true fishes. It is quite often regarded as
swim bladder or air bladder and found to be highly developed in Acanthopterygii (spiny rayed
teleosts).

It is an accessory respiratory organ, which also helps in sound production and sound perception,
fat storage (e.g., in the gonostomatid species). It is an important hydrostatic organ, which
contains a gas secreting complex, is composed of a gas gland covered with blood vessels.

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Respiration is supplemented by the gas bladder in many physostomous fishes with an open duct.
The gas bladder has undergone several modifications in various species of bony fishes (Fig. 5.13
a to f).
In chondrostei fishes such as Polypterus, the gas bladder is in the form of an unequal bilohed
structure with a small left lobe and large right lobe communicating with ventral part of the
pharynx (Fig. 5.13a). Both the lobes join together to a small opening called glottis provided
with a muscular sphincter. However, the Acipenser comprises oval shaped bladder of wide
opening into the oesophagus (Fig. 5.13b).

The holostean fishes like Lepidosteus has an unpaired sac which opens into the oesophagus by a
glottis. (Fig. 5.13c). The wall of bladder is composed of the fibrous bands produced in the alveoli
arranged into two rows. Each alveoli is further subdivided into smaller sacculi.

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In Amia, the gas bladder is very large and its wall is highly sacculated. These fishes can survive
in water depleted with oxygen, if they are capable to swallow air, which then passes into the gas
bladder through a pneumatic duct.
In Amia the gas bladder is relatively important as it lives in the temperate regions of North
America. This fish frequently ascends for air when the temperature of well aerated water
increases to 25C.

Since the gas bladder of physostomous fishes contains more carbon dioxide than the atmospheric
air, it has been considered that removal of this waste gas is performed there also. The dipnoi
fishes possess a well-developed gas bladder which is structurally similar to the amphibian lungs.

The gas bladder is a large unpaired sac-like in Neoceratodus which contains one dorsal and one
ventral fibrous ridges projecting in to this cavity (Fig. 5.13b).

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Many alveoli are formed due to the presence of transverse septa in between these ridges. The
alveoli in turn are further subdivided into several smaller sacculi. The complexities in the gas
bladder increase in the Protopterus and Lepidosiren which have lung-like bladder. (Fig. 5.13e).

The gas bladder is present in many teleosts while in others it is completely absent such as in
Echeneiformes, Symbranchiformes, Saccopharyngiformes and Gobeisociformes. If present, the
gas bladder may be oval, fusiform, tubular, heart shaped, horse-shoe shaped or dumb-bell
shaped.

In Cyprinidae the gas bladder lies freely in the abdominal cavity or may be attached to the
vertebral column by fibrous tissue. It has two chambers, interconnected with each other (Fig.
5.13 f).

The members of Sparidae, Notopteridae and Scombridae possess paired caeca4ike gas bladder
extended into the tail. In some fishes, like Clarias batrachus and Heteropneustes fossilis, the gas
bladder is reduced and lies enclosed in bone.

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The fishes living in torrential waters of the hills have contain rudimentary gas bladder having
only small anterior lobe enclosed in bone and no posterior lobe (Psillorhynchus and
Nemacheilus).

In most of the sound producing fishes, the gas bladder is provided with caecal outgrowths. In
Gadus a pair of caecal outgrowths arises from the gas bladder and projects in the head region
while in Otolithus each anterolateral side of the gas baldder gives off a caecal outgrowth which
immediately divide into two branches.

One branch runs anteriorly while another goes to the posterior. The caeca are much branched in
Corviva lobata and arises from whole periphery of the gas bladder.
The gas bladder is rarely divided completely by the septum. Most often it is partially subdivided
by incomplete septum. All teleosts in the beginning generally have an open duct of gas bladder,
i.e., they are physostomous but in later stages it closes in many teleosts and they become
physoclistic (Fig. 5.14).

2. Blood Supply of Gas Bladder:

The gas bladder is supplied with blood from the posterior branches of dorsal aorta or from
coeliacomesenteric artery. In some fishes the venous blood is collected by a vessels of hepatic
portal system while in others the gas bladder vein collects the venous blood and discharges it into
posterior cardinal vein.

The vascularization of gas bladder differs from species to species. In physostomous carps, the
inner surface of the bladder is covered at frequent places by blood vessels arranged in a fan-like
manner. These vessels form red patches of various shapes and sizes, are known as red bodies
which is a countercurrent arrangement of small arterioles and venules that constitute a rete
mirabile (Fig. 5.15a, b).
Before entering the tissue, artery divides into a large number of small capillaries, they are
parallel to a series of venous capillaries leaving the tissue.

The arterial capillaries are surrounded by venous capillaries and vice versa, forming an
extensive exchange surface between inflowing and outflowing blood. Retail capillaries serve to
transfer heat or gases between arterial blood entering the tissue and venous blood leaving it.

In physostomous fishes, the rete mirabile is rather primitive and is covered with flattened
epithelium, known as red bodies, while in physoditus fishes the capillaries are covered by thick
glandular folded epithelium and is called as red gland. In some fishes like Clupidae and
Salmonidae the blood vessels are uniformly distributed over the bladder and do not form a rete
mirabile.

3. Histology of Gas Bladder:

In cyprinids the anterior chamber of gas bladder comprises.

1. An innermost epithelial layer.

2. Lamina propria of thin connective tissue layer.

3. Muscularis mucosa of thick layer of smooth muscle fibres.

4. Submucosa of loose connective tissue.

5. An outermost tunica externa of dense collegenous muscle fibres.

However, posterior chamber of the gas bladder differs histologically, and comprises of a
glandular layer of large cells containing fine granulated cytoplasm which lie inside the tunica
externa. The glandular part of gas bladder is richly supplied by blood capillaries. The muscles of
posterior chamber is also known to have regulatory function of gas gland and to control the
volume of gas bladder.

In some fishes the anterior chamber of gas bladder contains a gas gland, which secretes gas
whereas the posterior chamber is thin-walled and helps in gas diffusion as in Synganthidae
species. In these fishes the gas bladder is closed and partially subdivided into two chambers.

However, in the Cyprinids it has pneumatic duct and gas gland is present in the posterior
chamber, which performs hydrostatic function, while the anterior chamber plays auditory
function (Fig. 5.16).

4. Gas Bladder in Sound Production:

Various branches arising from the vagus nerve and from the coeliac ganglia innervate the gas
bladder. These nerves terminate in re-absorbent area, the oval, the rete and in the secondary
epithelium. The muscular wall of the bladder is also very well supplied with nerves. Out of
twenty thousand fish species only few hundred species are known to produce sound of various
intensities.

In fishes generally three sonic mechanisms work for sound production:

i. Hydrodynamic:

Sound produced as a result of swimming movements particularly when rapid changes in

direction or velocity occurs.

ii. Stridulatory:

Sound produced by rubbing of teeth, fin spines and bones. Ex. grunts, pomadasyidae.

iii. By Gas Bladder:

Sound is produced by vibrations of striated muscle, which originates from dorsal body wall and
inserts on the gas bladder. Ex. grenadieres (melanonidae), drums (Sciaenidae). Toad fishes are
able to produce sound by rapid change in the volume of the gas bladder.

Sound produced by the gas bladder usually has low pitches, however, sound made by teeth or
bones has higher frequencies. Sound plays important roles in breeding behaviour and in the
defence as well.

5. Gas Bladder in Sound Reception:

The sound waves easily pass from the sea water to fish body because of similar densities. But
these sound waves are discontinued by the gas bladder and therefore, the gas bladder acts as a
sound conductors or resonator.

In fishes like cods (Gadidae) and orgies (Sparidae) the gas bladder is extended in such a way that
it touches the bones near the sacculus of the internal ear, the variation in pressure due to sound
waves may be transmitted directly to the perilymph.

Extension of the gas bladder grows in the form of cartilaginous capsule, i.e., prootic and pterotic
bullae, lies closely to the perilymph spaces of superior and inferior part of the internal ear.

In the order Cypriniformes the gas bladder transmits the sound waves to the internal ear by as
special apparatus which consists of series of paired bones or ossicles and is known a Weberian
apparatus, which connects the gas bladder to the internal ear. These ossicles originate from the
apophysis of anterior vertebrae.

Weberian apparatus consists of five ossicles, i.e., claustrum, scaphium, intercalarium and the
tripus, which do not show homology with the mammalian ear, hence called Weberian ossicles.
The posterior most ossicle is tripus, which is largest and triangular piece.

Posteriorly it touches anterior wall of the gas bladder while anteriorly it articulates to the
ligaments of next bone, i.e., intercalar. But when the latter is absent it is attached to scaphium
which in turn is attached to the minute anterior most claustrum.
The claustrum touches a membranes atrium sinus impar, that lies in the basioccipetal bone of the
head and is an extension of the perilymph system of the internal ear. In Gymnotids
(Gymnotidae), the scaphium touches the atrium sinus impar due to absence of claustrum. The
intercalarium also shows variations in the structure and mode of its development.

It may be a small nodule like bone in the ligament, separated from the vertebral column as found
in the siluroids (Siluridae). Sometimes it may develop as rod-like extension touching the centrum
of second vertebra as in the carp (Abeo, Cirrhina and Tor).

The Weberian ossicles provide a connection between the gas bladder and the internal ear by a
series, i.e., gas bladder Weberian ossicle sinus impar sinus endolymphaticus
transverse canal sacculus.

At the time of functioning of Weberian ossicles, volume of gas bladder changes due to which the
gas bladder moves in such a manner that pressure changes are transmitted to the perilymph and
hence to the sensory cells of inferior part of the labyrinth which is the seat of sound reception.

In some species the gas bladder is enclosed in a bony capsule or connective tissue and projects
through a small aperture to attach the tripus. A change in the volume of gas bladder due to its
rhythmic compression causes its wall to bulge out and push the ossicles forwards.

Among the Cypriniformes fishes wide range of sound perception and better sound discrimination
are seen than those fishes which do not possess Weberian apparatus. Removal of gas bladder in
fish-like minnows, greatly reduces the auditory range.

6. Gas Bladder as a Hydrostatic Organs:

The density of fish flesh is greater than that of water. To make the body weightless and to
minimize the energy consumption in maintaining the body position, the fish stores fats and oils
in muscle and liver, fills the oxygen in the gas bladder. In this way fish can reduce its body
weight.

In bony fishes the gas bladder brings the density of fish closely to that of surrounding water. In
sharks and rays the air bladder is absent and they maintain their body buoyancy by regulating the
water ballast present in the body cavity and operated through their abdominal pores.

In marine fishes the gas bladder can make up 4 to 11 percent of the body volume while in
freshwater fishes 7 to 11 percent of the body volume is maintained by the gas bladder.

The fishes may be divided into physostomous (bladder with opening into gut) an physoclitous
(bladder closed) on the basis of its functional and morphological differences. The change of one
condition to another is a gradual process and is concerned with gas secreting and resorbing
structures.
In many physostomous species the gas bladder loses the pneumatic duct which was open outside
in the young. The condition is known as paraphysoclistious as found in lantern fishes
(Myctophidae).

The soft-rayed fishes (Malacoptergii) are physostomous and the spiny rayed ones
(Acanthopterygii) are physoclitous. In true physoclistous teleosts the pressure in the gas bladder
is adjusted through secretion or resorption of gasses from or to the blood.

The position of the gas bladder in relation to the centre of gravity of fish plays an important role
in swimming and maintaining its position. Normal swimming position of fish is maintained
effortlessly with the help of gas bladder. Some fishes can displace their gas bladder to achieve
their normal position from unusual upside down swimming position of the body.

7. Filling and Emptying of the Gas Bladder:

Gas bladder has unique character that it stores 500 times oxygen and 30 times nitrogen.
Physostomous fishes such as trouts and salmon fill their gas bladder by gulping air at the time of
removal of yolk sac. Although adults of these fishes are able to secrete and absorb gas through
the blood supply but at initial stage they have to depend on atmosphere to fill their gas bladder.

Many physoclistous fishes like sticklebacks (Gastrosteus), guppy (Lebistes) and seahorse
(Hippocampus) possess pneumatic duct in larval stage, thus first filling of the gas bladder takes
place from atmospheric air.

Some deep sea fishes such as grenadiers (Melanonidae) have functional gas bladder with
different mechanism for initial filling of gas bladder unless they are pelagic in early life stages.
Fishes are able to change gas content in such a way that the gas volume is almost constant
regardless of hydrostatic pressure. Boyles law which states that the volume of gas changes
inversely with pressure, is also applicable to the gas bladder.

8. Secretion of Gas from Blood to Lumen of Bladder:

The gases contained in the blood is released into the cavity of the gas bladder through highly
vascular regions, called gas secreting complex present in the wall of the bladder. The gas
secreting complex consists of (i) gas gland and (ii) rete mirabile.

The gas gland is the region of the bladder epithelium and may be one layered, folded or made up
to multilayered stratified epithelium. Rete miraibile is small blood vessels underlying the
epithelium.

Arteries and veins of the bladder make intimate diffusional contact with each other and forms
countercurrent multiplier system which ensures concentration difference of many substances
from one end to another end of the organ (Fig. 15). Deep sea fishes such as searobins (Trigla)
usually fill their gas bladder with oxygen.

9. Reabsorption of Gas from Bladder:

It is accomplished as follows:

1. The gas from bladder may be diffused into the blood vessels present throughout the gas
bladder wall, other than the gas secreting complex as found in killifishes (Cyprinodontidae) and
sauries (Scombresocidae).

2. Generally the gas is drained from single chambered or posterior sac of gas bladder through a
thin area of the bladder wall which comprises a network of capillaries separated from the lumen
of bladder through a very thin area contained in capillaries in the bladder wall is known as oval
organ.

Sphincter surrounds the oval organ and regulates that rate of gas reabsorption by dilating and
contracting the oval orifice, e.g, cods and spiny rayed fishes, i.e., Acanthopterygii.