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

HOW ARE GILLS EFFICIENT? 1) A short diffusion. 3) Gills have little physiological dead space. 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. the blood in the filament folds travels forward. 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. A single gill of a bony fish consists of a curved gill arch bearing a V-shaped double row of gill filaments. giving it a sort of fuzzy appearance and increasing the amount of surface area along a given length of filament. so that a constant imbalance is maintained between the lower amount of oxygen in the blood and the higher amount in the water. Each filament has many minute folds in its surface.000 to 1/10. ensuring passage of oxygen to the blood.extract as much as 80% of the oxygen contained in the water passing over the gills. . the surface area of the gills is commonly 10 to 60 times more than that of the whole body surface. 2) By using countercurrent circulation in the gill. Consequently. being separated from it by a very thin membrane usually 1 to 3 microns (4/100. 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. or travel. whereas humans can extract only about 25% of the oxygen from the air taken into the lungs. If the blood were to flow in the same direction as the water. The oxygenated water flows through the tinny gill filaments and it exchanges the carried oxygen for carbon dioxide through a process called diffusion. distance for the oxygen increases the rate of oxygen entry into the blood.000 in) or less thick. in the opposite direction to the water flow. The blood traveling in the folds of the filaments is very close to the oxygen- containing water.

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. 4) Water flows continuously in only one direction over the gills. two-way flow of air in and out of lungs of mammals. as contrasted with the interrupted. IN CONCLUSION. ...

This chapter focuses on the structure of fish air-breathing organs (ABO). Although the pattern of reduced Hb-O2 affinity is seen among closely related species of non-air-breathing and air-breathing osteoglossids and erythrinids. Few structures other than the mouth and jaws can be used to capture air. and absorption of aerial oxygen. the form taken by the ABOs of modern fishes is characterized by either evolutionary parallelism or convergence driven by limitations of structure. their air-ventilation mechanics. A variety of factors. in rocky intertidal zones. and background knowledge required to evaluate the presence of air breathing in a species. and the timing and sequence of air inhalation and expiration. and surface area. Thus. retention. This chapter discusses in detail the comparative morphometrics of ABOs. and in marshes and bays. and examines the habitats of extant air-breathing fishes. Marine air breathers occur on tropical coral reefs. The principal requirements for an ABO are the acquisition. The chapter also presents a simplified ABO classification scheme that groups all the primitive fishes possessing lungs or respiratory gas bladders. 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. This chapter reviews physical principles governing the suitability of air and water as respiratory media. and comparative physiology. This chapter examines the influence of aerial gas exchange on the blood-respiratory properties of air breathing fishes. fishes have a tetrameric Hb contained within a nucleated erythrocyte. Research indicates that the degree of branchial isolation enjoyed by a species. In terms of holding inspired air. branchial region. The chapter also presents a general classification of the types of air breathing fishes. The chapter discusses the gill and skin morphology in relation to air breathing. comparisons among a diversity of air-breathing and non-air-breathing species fail to verify this as a universal evolutionary pattern. The chapter provides an operational definition of fish air breathing along with a discussion of the types of observations. vertebrate evolution. With few exceptions. space.This chapter provides a general overview of fish air breathing. 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. to the periodic aerial exposure imposed by low tides. 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. Air breathing has persisted throughout the evolutionary history of the fishes and has played a fundamental role in this group's evolution. from the adoption of an amphibious behavior to exploit resources at the air–water interface. or digestive track. experimental evidence. Modern air-breathing fishes occur in a variety of freshwater and marine habitats. tidal volumes. that . A high [Hb] can serve both an O2 storage and a blood- buffering function in air-breathing fishes. and even aquatic hypoxia in certain habitats have all played a role in selection for air breathing among marine species. The chapter also defines terms and concepts related to aerial and aquatic gas exchange and bimodal respiration.

Blood Supply of Gas Bladder 3. Filling and Emptying of the Gas Bladder 8. Gas Bladder in Sound Production 5. Gas Bladder as a Respiratory Device 2. fish may ascend for air as frequently as 1200 times in 24 hours. For amphibious air breathers. and elucidated respiratory organ function during bimodal respiration. determined respiratory capacity. is a major determinant of Hb-O2 affinity during air breathing. Gas Bladder as a Hydrostatic Organs 7. In situations of chronic hypoxia. The chapter emphasizes the comparative physiological aspects of air breathing. Secretion of Gas from Blood to Lumen of Bladder 9. Gas Bladder in Sound Production 5. its degree of independence from gill respiration. Reabsorption of Gas from Bladder. Histology 4. Gas Bladder as a Respiratory Device 2. Gas Bladder in Fishes (With Diagram) Article shared by In this article we will discuss about:. This chapter begins with a historical overview. Gas Bladder in Sound Reception . For most aquatic air breathers. Blood Supply of Gas Bladder 3. Light level also affects air-breath duration as can the presence of water-borne toxins. ].1. Air-breath durations differ among species. followed by a comparison of air breathing cycles and a review of whole-organism respiratory gas exchange. aerial respiration is utilized to the extent needed to supplement aquatic respiration and sustain normal VO2.is. These are highly variable. Aerial and Aquatic Gas Exchange This chapter reviews the diverse information that is found under the general subject of “air-breathing fish respiration. Gas Bladder in Sound Reception 6. the complete transition from aquatic to aerial gas exchange ensures a period of total respiratory independence from water. The chapter also examines the specialized aspects of respiratory organ function. and are affected by several physical factors including aquatic oxygen level and temperature. An effect of aquatic CO2 has also been suggested for some species. Contents: 1. Histology of Gas Bladder 4.” Studies of air-breathing fish respiration have identified factors regulating the duration of the air-breath cycle.

6. which also helps in sound production and sound perception. Reabsorption of Gas from Bladder 1. It is an important hydrostatic organ. Gas Bladder as a Hydrostatic Organs 7. 5. Secretion of Gas from Blood to Lumen of Bladder 9. Filling and Emptying of the Gas Bladder 8. 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). . which contains a gas secreting complex.13 a to f).g. It is an accessory respiratory organ. fat storage (e. in the gonostomatid species). ADVERTISEMENTS: Respiration is supplemented by the gas bladder in many physostomous fishes with an open duct. is composed of a gas gland covered with blood vessels.. The gas bladder has undergone several modifications in various species of bony fishes (Fig.

13b).13a). . Both the lobes join together to a small opening called ‘glottis’ provided with a muscular sphincter. However. the gas bladder is very large and its wall is highly sacculated. ADVERTISEMENTS: In Amia. 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.13c). Each alveoli is further subdivided into smaller sacculi. 5.In chondrostei fishes such as Polypterus. These fishes can survive in water depleted with oxygen. which then passes into the gas bladder through a pneumatic duct. if they are capable to swallow air. The holostean fishes like Lepidosteus has an unpaired sac which opens into the oesophagus by a glottis. The wall of bladder is composed of the fibrous bands produced in the alveoli arranged into two rows. 5. (Fig. 5. the Acipenser comprises oval shaped bladder of wide opening into the oesophagus (Fig.

heart shaped. . (Fig.In Amia the gas bladder is relatively important as it lives in the temperate regions of North America. the gas bladder is reduced and lies enclosed in bone. In some fishes. If present. 5. 5. horse-shoe shaped or dumb-bell shaped. 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. In most of the sound producing fishes. ADVERTISEMENTS: 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). One branch runs anteriorly while another goes to the posterior. it has been considered that removal of this waste gas is performed there also. the gas bladder is provided with caecal outgrowths. Symbranchiformes. Saccopharyngiformes and Gobeisociformes. The gas bladder is present in many teleosts while in others it is completely absent such as in Echeneiformes. This fish frequently ascends for air when the temperature of well aerated water increases to 25°C. Since the gas bladder of physostomous fishes contains more carbon dioxide than the atmospheric air. It has two chambers. The dipnoi fishes possess a well-developed gas bladder which is structurally similar to the amphibian lungs. ADVERTISEMENTS: Many alveoli are formed due to the presence of transverse septa in between these ridges. tubular. The members of Sparidae. fusiform. The complexities in the gas bladder increase in the Protopterus and Lepidosiren which have lung-like bladder. like Clarias batrachus and Heteropneustes fossilis.13b). In Cyprinidae the gas bladder lies freely in the abdominal cavity or may be attached to the vertebral column by fibrous tissue.13 f). interconnected with each other (Fig. The caeca are much branched in Corviva lobata and arises from whole periphery of the gas bladder.13e). 5. Notopteridae and Scombridae possess paired caeca4ike gas bladder extended into the tail. The alveoli in turn are further subdivided into several smaller sacculi. the gas bladder may be oval. 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.

In physostomous carps. Most often it is partially subdivided by incomplete septum. . These vessels form red patches of various shapes and sizes. 5. b).15a.14). they are physostomous but in later stages it closes in many teleosts and they become physoclistic (Fig. All teleosts in the beginning generally have an open duct of gas bladder. are known as ‘red bodies’ which is a countercurrent arrangement of small arterioles and venules that constitute a ‘rete mirabile’ (Fig.. i. 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.e. Blood Supply of Gas Bladder: The gas bladder is supplied with blood from the posterior branches of dorsal aorta or from coeliacomesenteric artery. The vascularization of gas bladder differs from species to species.The gas bladder is rarely divided completely by the septum. 2. the inner surface of the bladder is covered at frequent places by blood vessels arranged in a fan-like manner. 5.

. Lamina propria of thin connective tissue layer. 4. artery divides into a large number of small capillaries. 3. The ‘arterial’ capillaries are surrounded by ‘venous’ capillaries and vice versa. 2. Retail capillaries serve to transfer heat or gases between arterial blood entering the tissue and venous blood leaving it. the rete mirabile is rather primitive and is covered with flattened epithelium. known as ‘red bodies’. Histology of Gas Bladder: In cyprinids the anterior chamber of gas bladder comprises. In some fishes like Clupidae and Salmonidae the blood vessels are uniformly distributed over the bladder and do not form a rete mirabile. 1.Before entering the tissue. In physostomous fishes. forming an extensive exchange surface between inflowing and outflowing blood. An outermost tunica externa of dense collegenous muscle fibres. An innermost epithelial layer. 3. Muscularis mucosa of thick layer of smooth muscle fibres. Submucosa of loose connective tissue. while in physoditus fishes the capillaries are covered by thick glandular folded epithelium and is called as ‘red gland’. 5. they are parallel to a series of venous capillaries leaving the tissue.

4. The muscular wall of the bladder is also very well supplied with nerves. posterior chamber of the gas bladder differs histologically. which secretes gas whereas the posterior chamber is thin-walled and helps in gas diffusion as in Synganthidae species. and comprises of a glandular layer of large cells containing fine granulated cytoplasm which lie inside the tunica externa. the rete and in the secondary epithelium. Gas Bladder in Sound Production: Various branches arising from the vagus nerve and from the coeliac ganglia innervate the gas bladder. while the anterior chamber plays auditory function (Fig. These nerves terminate in re-absorbent area. The glandular part of gas bladder is richly supplied by blood capillaries. In these fishes the gas bladder is closed and partially subdivided into two chambers.16). However. In some fishes the anterior chamber of gas bladder contains a gas gland. the oval.However. Out of twenty thousand fish species only few hundred species are known to produce sound of various intensities. 5. In fishes generally three sonic mechanisms work for sound production: . in the Cyprinids it has pneumatic duct and gas gland is present in the posterior chamber. which performs hydrostatic function. The muscles of posterior chamber is also known to have regulatory function of gas gland and to control the volume of gas bladder.

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. Sound plays important roles in breeding behaviour and in the defence as well. Ex. intercalarium and the tripus. which connects the gas bladder to the internal ear. prootic and pterotic bullae. however.. hence called ‘Weberian ossicles’.i. the gas bladder acts as a sound conductors or resonator. 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. pomadasyidae.e. These ossicles originate from the apophysis of anterior vertebrae. Stridulatory: Sound produced by rubbing of teeth. 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. Ex. iii. The posterior most ossicle is tripus. Extension of the gas bladder grows in the form of cartilaginous capsule. i. grunts. which originates from dorsal body wall and inserts on the gas bladder. intercalar.. scaphium. Posteriorly it touches anterior wall of the gas bladder while anteriorly it articulates to the ligaments of next bone.. By Gas Bladder: Sound is produced by vibrations of striated muscle. ii. drums (Sciaenidae).e. grenadieres (melanonidae). Weberian apparatus consists of five ossicles. Hydrodynamic: Sound produced as a result of swimming movements particularly when rapid changes in direction or velocity occurs. claustrum. . 5. the variation in pressure due to sound waves may be transmitted directly to the perilymph. which do not show homology with the mammalian ear.e. fin spines and bones. lies closely to the perilymph spaces of superior and inferior part of the internal ear. i. sound made by teeth or bones has higher frequencies. Gas Bladder in Sound Reception: The sound waves easily pass from the sea water to fish body because of similar densities. But when the latter is absent it is attached to scaphium which in turn is attached to the minute anterior most claustrum. i. which is largest and triangular piece. But these sound waves are discontinued by the gas bladder and therefore.

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. fills the oxygen in the gas bladder. the fish stores fats and oils in muscle and liver. 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 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.e. the scaphium touches the atrium sinus impar due to absence of claustrum. separated from the vertebral column as found in the siluroids (Siluridae). greatly reduces the auditory range. To make the body weightless and to minimize the energy consumption in maintaining the body position. 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. Among the Cypriniformes fishes wide range of sound perception and better sound discrimination are seen than those fishes which do not possess Weberian apparatus.The claustrum touches a membranes atrium sinus impar. In Gymnotids (Gymnotidae).. 6. Removal of gas bladder in fish-like minnows. The intercalarium also shows variations in the structure and mode of its development. 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. A change in the volume of gas bladder due to its rhythmic compression causes its wall to bulge out and push the ossicles forwards. In bony fishes the gas bladder brings the density of fish closely to that of surrounding water. In this way fish can reduce its body weight. i. Cirrhina and Tor). Gas Bladder as a Hydrostatic Organs: The density of fish flesh is greater than that of water. It may be a small nodule like bone in the ligament. At the time of functioning of Weberian ossicles. Sometimes it may develop as rod-like extension touching the centrum of second vertebra as in the carp (Abeo. The change of one condition to another is a gradual process and is concerned with gas secreting and resorbing structures. gas bladder → Weberian ossicle → sinus impar → sinus endolymphaticus → transverse canal → sacculus. The Weberian ossicles provide a connection between the gas bladder and the internal ear by a series. that lies in the basioccipetal bone of the head and is an extension of the perilymph system of the internal ear.

The condition is known as paraphysoclistious as found in lantern fishes (Myctophidae). 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. thus first filling of the gas bladder takes place from atmospheric air. 8. The soft-rayed fishes (Malacoptergii) are physostomous and the spiny rayed ones (Acanthopterygii) are physoclitous. Physostomous fishes such as trouts and salmon fill their gas bladder by gulping air at the time of removal of yolk sac. The ‘gas secreting complex’ consists of (i) gas gland and (ii) rete mirabile. Normal swimming position of fish is maintained effortlessly with the help of gas bladder. called ‘gas secreting complex’ present in the wall of the bladder. folded or made up to multilayered stratified epithelium. is also applicable to the gas bladder. Boyle’s law which states that the volume of gas changes inversely with pressure. The gas gland is the region of the bladder epithelium and may be one layered.In many physostomous species the gas bladder loses the pneumatic duct which was open outside in the young. 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 . 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. guppy (Lebistes) and seahorse (Hippocampus) possess pneumatic duct in larval stage. Rete miraibile is small blood vessels underlying the epithelium. 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. 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). 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. Fishes are able to change gas content in such a way that the gas volume is almost constant regardless of hydrostatic pressure. In true physoclistous teleosts the pressure in the gas bladder is adjusted through secretion or resorption of gasses from or to the blood.

15). i. 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. Acanthopterygii. 2.g.. other than the gas secreting complex as found in killifishes (Cyprinodontidae) and sauries (Scombresocidae). 9.from one end to another end of the organ (Fig.e. Reabsorption of Gas from Bladder: It is accomplished as follows: 1. cods and spiny rayed fishes. The gas from bladder may be diffused into the blood vessels present throughout the gas bladder wall. e. Sphincter surrounds the oval organ and regulates that rate of gas reabsorption by dilating and contracting the oval orifice. Deep sea fishes such as searobins (Trigla) usually fill their gas bladder with oxygen. .