Swim bladder

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The swim bladder of a rudd Swim bladder.jpg
The swim bladder of a rudd
Internal positioning of the swim bladder of a bleak
S: anterior, S': posterior portion of the air bladder
oe: oesophagus; l: air passage of the air bladder Air bladder in a bleak.jpg
Internal positioning of the swim bladder of a bleak
S: anterior, S': posterior portion of the air bladder
œ: œsophagus; l: air passage of the air bladder

The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish [1] ) to control their buoyancy, and thus to stay at their current water depth without having to waste energy in swimming. [2] Also, the dorsal position of the swim bladder means the center of mass is below the center of volume, allowing it to act as a stabilizing agent. Additionally, the swim bladder functions as a resonating chamber, to produce or receive sound.

Organ (anatomy) Collection of tissues

Organs are groups of tissues with similar functions. Plant and animal life relies on many organs that coexist in organ systems.

Buoyancy An upward force that opposes the weight of an object immersed in fluid

Buoyancy or upthrust, is an upward force exerted by a fluid that opposes the weight of an immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the pressure at the bottom of a column of fluid is greater than at the top of the column. Similarly, the pressure at the bottom of an object submerged in a fluid is greater than at the top of the object. The pressure difference results in a net upward force on the object. The magnitude of the force is proportional to the pressure difference, and is equivalent to the weight of the fluid that would otherwise occupy the volume of the object, i.e. the displaced fluid.

Energy Physical property transferred to objects to perform heating or work

In physics, energy is the quantitative property that must be transferred to an object in order to perform work on, or to heat, the object. Energy is a conserved quantity; the law of conservation of energy states that energy can be converted in form, but not created or destroyed. The SI unit of energy is the joule, which is the energy transferred to an object by the work of moving it a distance of 1 metre against a force of 1 newton.


The swim bladder is evolutionarily homologous to the lungs. Charles Darwin remarked upon this in On the Origin of Species . [3] Darwin reasoned that the lung in air-breathing vertebrates had derived from a more primitive swim bladder.

Homology (biology) existence of shared ancestry between a pair of structures, or genes, in different taxa

In biology, homology is the existence of shared ancestry between a pair of structures, or genes, in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats, the arms of primates, the front flippers of whales and the forelegs of dogs and horses are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.

Lung Essential respiration organ in many air-breathing animals

The lungs are the primary organs of the respiratory system in humans and many other animals including a few fish and some snails. In mammals and most other vertebrates, two lungs are located near the backbone on either side of the heart. Their function in the respiratory system is to extract oxygen from the atmosphere and transfer it into the bloodstream, and to release carbon dioxide from the bloodstream into the atmosphere, in a process of gas exchange. Respiration is driven by different muscular systems in different species. Mammals, reptiles and birds use their different muscles to support and foster breathing. In early tetrapods, air was driven into the lungs by the pharyngeal muscles via buccal pumping, a mechanism still seen in amphibians. In humans, the main muscle of respiration that drives breathing is the diaphragm. The lungs also provide airflow that makes vocal sounds including human speech possible.

Charles Darwin British naturalist, author of "On the origin of species, by means of natural selection"

Charles Robert Darwin, was an English naturalist, geologist and biologist, best known for his contributions to the science of evolution. His proposition that all species of life have descended over time from common ancestors is now widely accepted, and considered a foundational concept in science. In a joint publication with Alfred Russel Wallace, he introduced his scientific theory that this branching pattern of evolution resulted from a process that he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding.

In the embryonic stages, some species, such as redlip blenny, [4] have lost the swim bladder again, mostly bottom dwellers like the weather fish. Other fish—like the opah and the pomfret—use their pectoral fins to swim and balance the weight of the head to keep a horizontal position. The normally bottom dwelling sea robin can use their pectoral fins to produce lift while swimming.

<i>Ophioblennius atlanticus</i> species of fish

Ophioblennius atlanticus, also known as the redlip blenny and the horseface blenny, is a species of combtooth blenny, family Blenniidae, found primarily in the western central Atlantic ocean. Redlip blennies can be found in coral crests and shallow fringing reefs. They are highly territorial and attack intruders with two long, sharp canine teeth. The adults are found at depths of 10 to 20 meters, and the eggs are benthic. The adults may reach up to four inches in length when fully grown, and they have large reddish lips, from which they attained their names. Redlip blennies largely feed on algae.

Opah genus of fishes

Opahs are large, colorful, deep-bodied pelagic lampriform fishes comprising the small family Lampridae. Only two living species occur in a single genus: Lampris. One species is found in tropical to temperate waters of most oceans, while the other is limited to a circumglobal distribution in the Southern Ocean, with the 34°S as its northern limit. Two additional species, one in the genus Lampris and the other in the monotypic Megalampris, are only known from fossil remains. The extinct family, Turkmenidae, from the Paleogene of Central Asia, is closely related, though much smaller.

Pomfret family of fishes

Pomfrets are perciform fishes belonging to the family Bramidae. The family includes about 20 species.

The gas/tissue interface at the swim bladder produces a strong reflection of sound, which is used in sonar equipment to find fish.

Sonar technique that uses sound propagation

Sonar is a technique that uses sound propagation to navigate, communicate with or detect objects on or under the surface of the water, such as other vessels. Two types of technology share the name "sonar": passive sonar is essentially listening for the sound made by vessels; active sonar is emitting pulses of sounds and listening for echoes. Sonar may be used as a means of acoustic location and of measurement of the echo characteristics of "targets" in the water. Acoustic location in air was used before the introduction of radar. Sonar may also be used in air for robot navigation, and SODAR is used for atmospheric investigations. The term sonar is also used for the equipment used to generate and receive the sound. The acoustic frequencies used in sonar systems vary from very low (infrasonic) to extremely high (ultrasonic). The study of underwater sound is known as underwater acoustics or hydroacoustics.

Cartilaginous fish, such as sharks and rays, do not have swim bladders. Some of them can control their depth only by swimming (using dynamic lift); others store fats or oils with density less than that of seawater to produce a neutral or near neutral buoyancy, which does not change with depth.

Chondrichthyes class of fishes

Chondrichthyes is a class that contains the cartilaginous fishes: they are jawed vertebrates with paired fins, paired nares, scales, a heart with its chambers in series, and skeletons made of cartilage rather than bone. The class is divided into two subclasses: Elasmobranchii and Holocephali.

Structure and function

Swim bladder from a bony (teleost) fish Oste023c labelled.png
Swim bladder from a bony (teleost) fish
How gas is pumped into the swim bladder using counter-current exchange. GasbladderpumpingEng.png
How gas is pumped into the swim bladder using counter-current exchange.

The swim bladder normally consists of two gas-filled sacs located in the dorsal portion of the fish, although in a few primitive species, there is only a single sac. It has flexible walls that contract or expand according to the ambient pressure. The walls of the bladder contain very few blood vessels and are lined with guanine crystals, which make them impermeable to gases. By adjusting the gas pressurising organ using the gas gland or oval window the fish can obtain neutral buoyancy and ascend and descend to a large range of depths. Due to the dorsal position it gives the fish lateral stability.

Pressure Force distributed continuously over an area

Pressure is the force applied perpendicular to the surface of an object per unit area over which that force is distributed. Gauge pressure is the pressure relative to the ambient pressure.

Guanine Chemical compound of DNA and RNA

Guanine is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine. In DNA, guanine is paired with cytosine. The guanine nucleoside is called guanosine.

In physostomous swim bladders, a connection is retained between the swim bladder and the gut, the pneumatic duct, allowing the fish to fill up the swim bladder by "gulping" air. Excess gas can be removed in a similar manner.

In more derived varieties of fish (the physoclisti) the connection to the digestive tract is lost. In early life stages, these fish must rise to the surface to fill up their swim bladders; in later stages, the pneumatic duct disappears, and the gas gland has to introduce gas (usually oxygen) to the bladder to increase its volume and thus increase buoyancy. In order to introduce gas into the bladder, the gas gland excretes lactic acid and produces carbon dioxide. The resulting acidity causes the hemoglobin of the blood to lose its oxygen (Root effect) which then diffuses partly into the swim bladder. The blood flowing back to the body first enters a rete mirabile where virtually all the excess carbon dioxide and oxygen produced in the gas gland diffuses back to the arteries supplying the gas gland. Thus a very high gas pressure of oxygen can be obtained, which can even account for the presence of gas in the swim bladders of deep sea fish like the eel, requiring a pressure of hundreds of bars. [5] Elsewhere, at a similar structure known as the oval window, the bladder is in contact with blood and the oxygen can diffuse back out again. Together with oxygen, other gases are salted out[ clarification needed ] in the swim bladder which accounts for the high pressures of other gases as well. [6]

The combination of gases in the bladder varies. In shallow water fish, the ratios closely approximate that of the atmosphere, while deep sea fish tend to have higher percentages of oxygen. For instance, the eel Synaphobranchus has been observed to have 75.1% oxygen, 20.5% nitrogen, 3.1% carbon dioxide, and 0.4% argon in its swim bladder.

Physoclist swim bladders have one important disadvantage: they prohibit fast rising, as the bladder would burst. Physostomes can "burp" out gas, though this complicates the process of re-submergence.

The swim bladder in some species, mainly fresh water fishes (common carp, catfish, bowfin) is interconnected with the inner ear of the fish. They are connected by four bones called the Weberian ossicles from the Weberian apparatus. These bones can carry the vibrations to the saccule and the lagena (anatomy). They are suited for detecting sound and vibrations due to its low density in comparison to the density of the fish's body tissues. This increases the ability of sound detection. [7] The swim bladder can radiate the pressure of sound which help increase its sensitivity and expand its hearing. In some deep sea fishes like the Antimora , the swim bladder maybe also connected to the macula of saccule in order for the inner ear to receive a sensation from the sound pressure. [8] In red-bellied piranha, the swimbladder may play an important role in sound production as a resonator. The sounds created by piranhas are generated through rapid contractions of the sonic muscles and is associated with the swimbladder. [9]

Teleosts are thought to lack a sense of absolute hydrostatic pressure, which could be used to determine absolute depth. [10] However, it has been suggested that teleosts may be able to determine their depth by sensing the rate of change of swim-bladder volume. [11]


The West African lungfish possesses a lung homologous to swim bladders PSM V20 D769 Lepidosiren annectens using the air bladder as a lung.jpg
The West African lungfish possesses a lung homologous to swim bladders
The illustration of the swim bladder in fishes ... shows us clearly the highly important fact that an organ originally constructed for one purpose, namely, flotation, may be converted into one for a widely different purpose, namely, respiration. The swim bladder has, also, been worked in as an accessory to the auditory organs of certain fishes. All physiologists admit that the swimbladder is homologous, or “ideally similar” in position and structure with the lungs of the higher vertebrate animals: hence there is no reason to doubt that the swim bladder has actually been converted into lungs, or an organ used exclusively for respiration. According to this view it may be inferred that all vertebrate animals with true lungs are descended by ordinary generation from an ancient and unknown prototype, which was furnished with a floating apparatus or swim bladder.

Charles Darwin, 1859 [3]

Swim bladders are evolutionarily closely related (i.e., homologous) to lungs. Traditional wisdom has long held that the first lungs, simple sacs connected to the gut that allowed the organism to gulp air under oxygen-poor conditions, evolved into the lungs of today's terrestrial vertebrates and some fish (e.g., lungfish, gar, and bichir) and into the swim bladders of the ray-finned fish. In 1997, Farmer proposed that lungs evolved to supply the heart with oxygen. In fish, blood circulates from the gills to the skeletal muscle, and only then to the heart. During intense exercise, the oxygen in the blood gets used by the skeletal muscle before the blood reaches the heart. Primitive lungs gave an advantage by supplying the heart with oxygenated blood via the cardiac shunt. This theory is robustly supported by the fossil record, the ecology of extant air-breathing fishes, and the physiology of extant fishes. [12] In embryonal development, both lung and swim bladder originate as an outpocketing from the gut; in the case of swim bladders, this connection to the gut continues to exist as the pneumatic duct in the more "primitive" ray-finned fish, and is lost in some of the more derived teleost orders. There are no animals which have both lungs and a swim bladder.

The cartilaginous fish (e.g., sharks and rays) split from the other fishes about 420 million years ago, and lack both lungs and swim bladders, suggesting that these structures evolved after that split. [12] Correspondingly, these fish also have both heterocercal and stiff, wing-like pectoral fins which provide the necessary lift needed due to the lack of swim bladders. Teleost fish with swim bladders have neutral buoyancy, and have no need for this lift. [13]

Deep scattering layer

Most mesopelagic fishes are small filter feeders which ascend at night using their swimbladders to feed in the nutrient rich waters of the epipelagic zone. During the day, they return to the dark, cold, oxygen deficient waters of the mesopelagic where they are relatively safe from predators. Lanternfish account for as much as 65 percent of all deep sea fish biomass and are largely responsible for the deep scattering layer of the world's oceans. California headlightfish.png
Most mesopelagic fishes are small filter feeders which ascend at night using their swimbladders to feed in the nutrient rich waters of the epipelagic zone. During the day, they return to the dark, cold, oxygen deficient waters of the mesopelagic where they are relatively safe from predators. Lanternfish account for as much as 65 percent of all deep sea fish biomass and are largely responsible for the deep scattering layer of the world's oceans.

Sonar operators, using the newly developed sonar technology during World War II, were puzzled by what appeared to be a false sea floor 300–500 metres deep at day, and less deep at night. This turned out to be due to millions of marine organisms, most particularly small mesopelagic fish, with swimbladders that reflected the sonar. These organisms migrate up into shallower water at dusk to feed on plankton. The layer is deeper when the moon is out, and can become shallower when clouds obscure the moon. [14]

Most mesopelagic fish make daily vertical migrations, moving at night into the epipelagic zone, often following similar migrations of zooplankton, and returning to the depths for safety during the day. [15] [16] These vertical migrations often occur over large vertical distances, and are undertaken with the assistance of a swim bladder. The swim bladder is inflated when the fish wants to move up, and, given the high pressures in the mesoplegic zone, this requires significant energy. As the fish ascends, the pressure in the swimbladder must adjust to prevent it from bursting. When the fish wants to return to the depths, the swimbladder is deflated. [17] Some mesopelagic fishes make daily migrations through the thermocline, where the temperature changes between 10 and 20 °C, thus displaying considerable tolerance for temperature change.

Sampling via deep trawling indicates that lanternfish account for as much as 65% of all deep sea fish biomass. [18] Indeed, lanternfish are among the most widely distributed, populous, and diverse of all vertebrates, playing an important ecological role as prey for larger organisms. The estimated global biomass of lanternfish is 550–660 million metric tonnes, several times the entire world fisheries catch. Lanternfish also account for much of the biomass responsible for the deep scattering layer of the world's oceans. Sonar reflects off the millions of lanternfish swim bladders, giving the appearance of a false bottom. [19]

Human uses

In some Asian cultures, the swim bladders of certain large fishes are considered a food delicacy. In China they are known as fish maw, 花膠/鱼鳔, [20] and are served in soups or stews.

The vanity price of a vanishing kind of maw is behind the imminent extinction of the vaquita, the world's smallest dolphin species. Found only in Mexico's Gulf of California, the once numerous vaquita now number less than 60 in total. Vaquita die in gillnets [21] set to catch totoaba (the world's largest drum fish). Totoaba are being hunted to extinction for its maw, which can sell for as much $10,000 per kilogram.

Swim bladders are also used in the food industry as a source of collagen. They can be made into a strong, water-resistant glue, or used to make isinglass for the clarification of beer. [22] In earlier times they were used to make condoms. [23]

Swim bladder disease

Swim bladder disease is a common ailment in aquarium fish. A fish with swim bladder disorder can float nose down tail up, or can float to the top or sink to the bottom of the aquarium. [24]

Risk of injury

Many anthropogenic activities, like pile driving or even Seismic wave, that could result from climate change or natural causes, can create high-intensity sound waves that cause a certain amount of damage to fish that possess a gas bladder. Physostomes can release air in order to decrease the tension in the gas bladder that may cause internal injuries to other vital organs, while physoclisti can't expel air fast enough, making it more difficult to avoid any major injuries. [25] Some of the commonly seen injuries included ruptured gas bladder and renal Haemorrhage. These mostly affect the overall health of the fish and didn't affect their mortality rate. [25] Investigators used the High-Intensity-Controlled Impedance Fluid Filled (HICI-FT), a stainless-steel wave tube with an electromagnetic shaker. It simulates high-energy sound waves in aquatic far-field, plane-wave acoustic conditions. [26] [27]

Similar structures in other organisms

Siphonophores have a special swim bladder that allows the jellyfish-like colonies to float along the surface of the water while their tentacles trail below. This organ is unrelated to the one in fish. [28]

Related Research Articles

Osteichthyes superclass of fishes

Osteichthyes, popularly referred to as the bony fish, is a diverse taxonomic group of fish that have skeletons primarily composed of bone tissue, as opposed to cartilage. The vast majority of fish are members of Osteichthyes, which is an extremely diverse and abundant group consisting of 45 orders, and over 435 families and 28,000 species. It is the largest class of vertebrates in existence today. The group Osteichthyes is divided into the ray-finned fish (Actinopterygii) and lobe-finned fish (Sarcopterygii). The oldest known fossils of bony fish are about 420 million years old, which are also transitional fossils, showing a tooth pattern that is in between the tooth rows of sharks and bony fishes.

Deep sea fish

Deep-sea fish are fish that live in the darkness below the sunlit surface waters, that is below the epipelagic or photic zone of the sea. The lanternfish is, by far, the most common deep-sea fish. Other deep sea fishes include the flashlight fish, cookiecutter shark, bristlemouths, anglerfish, viperfish, and some species of eelpout.

Respiratory system A biological system of specific organs and structures for gas exchange in animals and plants

The respiratory system is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies greatly, depending on the size of the organism, the environment in which it lives and its evolutionary history. In land animals the respiratory surface is internalized as linings of the lungs. Gas exchange in the lungs occurs in millions of small air sacs called alveoli in mammals and reptiles, but atria in birds. These microscopic air sacs have a very rich blood supply, thus bringing the air into close contact with the blood. These air sacs communicate with the external environment via a system of airways, or hollow tubes, of which the largest is the trachea, which branches in the middle of the chest into the two main bronchi. These enter the lungs where they branch into progressively narrower secondary and tertiary bronchi that branch into numerous smaller tubes, the bronchioles. In birds the bronchioles are termed parabronchi. It is the bronchioles, or parabronchi that generally open into the microscopic alveoli in mammals and atria in birds. Air has to be pumped from the environment into the alveoli or atria by the process of breathing which involves the muscles of respiration.

Rete mirabile complex of arteries and veins lying very close to each other, found in some vertebrates, mainly warm-blooded ones.

A rete mirabile is a complex of arteries and veins lying very close to each other, found in some vertebrates, mainly warm-blooded ones. The rete mirabile utilizes countercurrent blood flow within the net to act as a countercurrent exchanger. It exchanges heat, ions, or gases between vessel walls so that the two bloodstreams within the rete maintain a gradient with respect to temperature, or concentration of gases or solutes. This term was coined by Galen.

Teleost infraclass of fishes

The teleosts or Teleostei are by far the largest infraclass in the class Actinopterygii, the ray-finned fishes, and make up 96% of all extant species of fish. Teleosts are arranged into about 40 orders and 448 families. Over 26,000 species have been described. Teleosts range from giant oarfish measuring 7.6 m (25 ft) or more, and ocean sunfish weighing over 2 t, to the minute male anglerfish Photocorynus spiniceps, just 6.2 mm (0.24 in) long. Including not only torpedo-shaped fish built for speed, teleosts can be flattened vertically or horizontally, be elongated cylinders or take specialised shapes as in anglerfish and seahorses. Teleosts dominate the seas from pole to pole and inhabit the ocean depths, estuaries, rivers, lakes and even swamps.

Fish anatomy study of the form or morphology of fishes

Fish anatomy is the study of the form or morphology of fishes. It can be contrasted with fish physiology, which is the study of how the component parts of fish function together in the living fish. In practice, fish anatomy and fish physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in living fish.

Euteleostomi clade

Euteleostomi is a successful clade that includes more than 90% of the living species of vertebrates. Euteleostomes are also known as "bony vertebrates". Both its major subgroups are successful today: Actinopterygii includes the majority of extant fish species, and Sarcopterygii includes the tetrapods.

Barracudina family of fishes

Barracudinas are any member of the marine mesopelagic fish family Paralepididae: 50 or so extant species are found almost worldwide in deep waters. Several genera are known only from fossils dating back to the Ypresian epoch.

Barotrauma Injury caused by pressure

Barotrauma is physical damage to body tissues caused by a difference in pressure between a gas space inside, or in contact with, the body, and the surrounding gas or fluid. The initial damage is usually due to over-stretching the tissues in tension or shear, either directly by expansion of the gas in the closed space or by pressure difference hydrostatically transmitted through the tissue. Tissue rupture may be complicated by the introduction of gas into the local tissue or circulation through the initial trauma site, which can cause blockage of circulation at distant sites or interfere with normal function of an organ by its presence.

Mesopelagic zone

The mesopelagiczone, also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins at the depth where only 1% of incident light reaches and ends where there is no light; the depths of this zone are between approximately 200 to 1000 meters below the ocean surface. It hosts a diverse biological community that includes bristlemouths, blobfish, bioluminescent jellyfish, giant squid, and a myriad of other unique organisms adapted to live in a low-light environment. It has long captivated the imagination of scientists, artists and writers; deep sea creatures are prominent in popular culture, particularly as horror movie villains.

Teleostomi Clade of jawed vertebrates

Teleostomi is an obsolete clade of jawed vertebrates that supposedly includes the tetrapods, bony fish, and the wholly extinct acanthodian fish. Key characters of this group include an operculum and a single pair of respiratory openings, features which were lost or modified in some later representatives. The teleostomes include all jawed vertebrates except the chondrichthyans and the extinct class Placodermi.

Lanternfish family of fishes

Lanternfishes are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represented by 246 species in 33 genera, and are found in oceans worldwide. They are aptly named after their conspicuous use of bioluminescence. Their sister family, the Neoscopelidae, are much fewer in number but superficially very similar; at least one neoscopelid shares the common name 'lanternfish': the large-scaled lantern fish, Neoscopelus macrolepidotus.

Pelagic fish Fish living in the pelagic zone of ocean or lake waters – being neither close to the bottom nor near the shore

Pelagic fish live in the pelagic zone of ocean or lake waters – being neither close to the bottom nor near the shore – in contrast with demersal fish, which do live on or near the bottom, and reef fish, which are associated with coral reefs.

Neutral buoyancy State of equilibrium between buoyancy and weight of a fully immersed object

Neutral buoyancy occurs when a object's average density is equal to the density of the fluid in which it is immersed, resulting in the buoyant force balancing the force of gravity that would otherwise cause the object to sink or rise. An object that has neutral buoyancy will neither sink nor rise.

Deep sea community Groups of organisms living deep below the sea surface sharing a habitat

A deep sea community is any community of organisms associated by a shared habitat in the deep sea. Deep sea communities remain largely unexplored, due to the technological and logistical challenges and expense involved in visiting this remote biome. Because of the unique challenges, it was long believed that little life existed in this hostile environment. Since the 19th century however, research has demonstrated that significant biodiversity exists in the deep sea.

Deep scattering layer A layer in the ocean consisting of a variety of marine animals that migrate vertically every day

The deep scattering layer, sometimes referred to as the sound scattering layer, is a name given to a layer in the ocean consisting of a variety of marine animals. It was discovered through the use of sonar, as ships found a layer that scattered the sound and was thus sometimes mistaken for the seabed. For this reason it is sometimes called the false bottom or phantom bottom. It can be seen to rise and fall each day in keeping with diel vertical migration.

Swim bladder disease

Swim bladder disease, also called swim bladder disorder or flipover, is a common ailment in aquarium fish. The swim bladder is an internal gas-filled organ that contributes to the ability of a fish to control its buoyancy, and thus to stay at the current water depth without having to waste energy in swimming. A fish with swim bladder disorder can float nose down tail up, or can float to the top or sink to the bottom of the aquarium.

Physoclisti are, collectively, fishes that lack a connection between the gas bladder and the alimentary canal, with the bladder serving only as a buoyancy organ.


Physostomes are fishes that have a pneumatic duct connecting the gas bladder to the alimentary canal. This allows the gas bladder to be filled or emptied via the mouth. This not only allows the fish to fill their bladder by gulping air, but also to rapidly ascend in the water without the bladder expanding to bursting point. In contrast, fish without any connection to their gas bladder are called physoclisti.

Fish physiology

Fish physiology is the scientific study of how the component parts of fish function together in the living fish. It can be contrasted with fish anatomy, which is the study of the form or morphology of fishes. In practice, fish anatomy and physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the later dealing with how those components function together in the living fish.


  1. "More on Morphology". www.ucmp.berkeley.edu.
  2. "Fish". Microsoft Encarta Encyclopedia Deluxe 1999. Microsoft. 1999.
  3. 1 2 Darwin, Charles (1859) Origin of Species Page 190, reprinted 1872 by D. Appleton.
  4. Nursall, J. R. (1989). "Buoyancy is provided by lipids of larval redlip blennies, Ophioblennius atlanticus". Copeia. 3 (3): 614–621. doi:10.2307/1445488. JSTOR   1445488.
  5. Pelster B (December 2001). "The generation of hyperbaric oxygen tensions in fish". News Physiol. Sci. 16 (6): 287–91. PMID   11719607.
  6. "Secretion Of Nitrogen Into The Swimbladder Of Fish. Ii. Molecular Mechanism. Secretion Of Noble Gases". Biolbull.org. 1981-12-01. Retrieved 2013-06-24.
  7. Kardong, Kenneth. Vertebrates: Comparative Anatomy, Function, Evolution. New York: McGraw-Hill Education. p. 701. ISBN   9780073524238.
  8. Deng, Xiaohong; Wagner, Hans-Joachim; Popper, Arthur N. (2011-01-01). "The inner ear and its coupling to the swim bladder in the deep-sea fish Antimora rostrata (Teleostei: Moridae)". Deep Sea Research Part I: Oceanographic Research Papers. 58 (1): 27–37. doi:10.1016/j.dsr.2010.11.001. PMC   3082141 . PMID   21532967.
  9. Onuki, A; Ohmori Y.; Somiya H. (January 2006). "Spinal Nerve Innervation to the Sonic Muscle and Sonic Motor Nucleus in Red Piranha, Pygocentrus nattereri (Characiformes, Ostariophysi)". Brain, Behavior and Evolution. 67 (2): 11–122. doi:10.1159/000089185. PMID   16254416.
  10. Bone, Q.; Moore, Richard H. Biology of fishes (3rd., Thoroughly updated and rev ed.). Taylor & Francis. ISBN   9780415375627.
  11. Taylor, Graham K.; Holbrook, Robert Iain; de Perera, Theresa Burt (6 September 2010). "Fractional rate of change of swim-bladder volume is reliably related to absolute depth during vertical displacements in teleost fish". Journal of the Royal Society Interface. 7 (50): 1379–1382. doi:10.1098/rsif.2009.0522. PMC   2894882 . PMID   20190038.
  12. 1 2 Farmer, Colleen (1997). "Did lungs and the intracardiac shunt evolve to oxygenate the heart in vertebrates" (PDF). Paleobiology. 23 (3): 358–372. doi:10.1017/S0094837300019734.
  13. Kardong, KV (1998) Vertebrates: Comparative Anatomy, Function, Evolution2nd edition, illustrated, revised. Published by WCB/McGraw-Hill, p. 12 ISBN   0-697-28654-1
  14. Ryan P "Deep-sea creatures: The mesopelagic zone" Te Ara - the Encyclopedia of New Zealand. Updated 21 September 2007.
  15. Moyle, Peter B.; Cech, Joseph J. (2004). Fishes : an introduction to ichthyology (5th ed.). Upper Saddle River, N.J.: Pearson/Prentice Hall. p. 585. ISBN   9780131008472.
  16. Bone, Quentin; Moore, Richard H. (2008). "Chapter 2.3. Marine habitats. Mesopelagic fishes". Biology of fishes (3rd ed.). New York: Taylor & Francis. p. 38. ISBN   9780203885222.
  17. Douglas EL, Friedl WA and Pickwell GV (1976) "Fishes in oxygen-minimum zones: blood oxygenation characteristics" Science, 191 (4230) 957–959.
  18. Hulley, P. Alexander (1998). Paxton, J.R.; Eschmeyer, W.N. (eds.). Encyclopedia of Fishes. San Diego: Academic Press. pp. 127–128. ISBN   978-0-12-547665-2.
  19. R. Cornejo; R. Koppelmann & T. Sutton. "Deep-sea fish diversity and ecology in the benthic boundary layer".
  20. Teresa M. (2009) A Tradition of Soup: Flavors from China's Pearl River Delta Page 70, North Atlantic Books. ISBN   9781556437656.
  21. "'Extinction Is Imminent': New report from Vaquita Recovery Team (CIRVA) is released". IUCN SSC - Cetacean Specialist Group. 2016-06-06. Retrieved 2017-01-25.
  22. Bridge, T. W. (1905) "The Natural History of Isinglass"
  23. Huxley, Julian (1957) "Material of early contraceptive sheaths." British Medical Journal, 1 (5018): 581–582.
  24. Johnson, Erik L. and Richard E. Hess (2006) Fancy Goldfish: A Complete Guide to Care and Collecting, Weatherhill, Shambhala Publications, Inc. ISBN   0-8348-0448-4
  25. 1 2 Halvorsen, Michele B.; Casper, Brandon M.; Matthews, Frazer; Carlson, Thomas J.; Popper, Arthur N. (2012-12-07). "Effects of exposure to pile-driving sounds on the lake sturgeon, Nile tilapia and hogchoker". Proceedings of the Royal Society B: Biological Sciences. 279 (1748): 4705–4714. doi:10.1098/rspb.2012.1544. ISSN   0962-8452. PMC   3497083 . PMID   23055066.
  26. Halvorsen, Michele B.; Casper, Brandon M.; Woodley, Christa M.; Carlson, Thomas J.; Popper, Arthur N. (2012-06-20). "Threshold for Onset of Injury in Chinook Salmon from Exposure to Impulsive Pile Driving Sounds". PLOS ONE. 7 (6): e38968. doi:10.1371/journal.pone.0038968. ISSN   1932-6203. PMC   3380060 . PMID   22745695.
  27. Popper, Arthur N.; Hawkins, Anthony (2012-01-26). The Effects of Noise on Aquatic Life. Springer Science & Business Media. ISBN   9781441973115.
  28. Clark, F. E.; C. E. Lane (1961). "Composition of float gases of Physalia physalis". Fed. Proc. 107 (3): 673–674. doi:10.3181/00379727-107-26724.

Further references