Swim bladder
The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled
The swim bladder is evolutionarily homologous to the lungs of tetrapods and lungfish. 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 as a specialized form of enteral respiration.
In the embryonic stages, some species, such as
The gas/tissue interface at the swim bladder produces a strong reflection of sound, which is used in sonar equipment to find fish.
Structure and function
The swim bladder normally consists of two gas-filled sacs located in the
In
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. This process begins with the acidification of the blood in the rete mirabile when the gas gland excretes lactic acid and produces carbon dioxide, the latter of which acidifies the blood via the bicarbonate buffer system. The resulting acidity causes the hemoglobin of the blood to lose its oxygen (Root effect) which then diffuses partly into the swim bladder. Before returning to the body, the blood re-enters the rete mirabile, and as a result, virtually all the excess carbon dioxide and oxygen produced in the gas gland diffuses back to the arteries supplying the gas gland via a countercurrent multiplication loop. 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
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 (
Evolution
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 vertebrateanimals: 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. The first lungs originated in the last common ancestor of the Actinopterygii (ray-finned fish) and Sarcopterygii (lobe-finned fish and the tetrapods) as expansions of the upper digestive tract which allowed them to gulp air under oxygen-poor conditions.[12] In the Actinopteri (ray-finned fish minus the bichirs) the lungs evolved into a swim bladder (secondary absent in some lineages), which unlike lungs that bud ventrally, buds dorsally from the anterior foregut.[13][14] Coelacanths have a "fatty organ" that have sometimes been referred to as a swim bladder, but is structurally different and have a separate evolutionary history.[15]
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.[16] 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.
As an adaptation to migrations between the surface and deeper waters, some fish have evolved a swim bladder where the gas is replaced with low-density wax esters as a way to cope with Boyle's law.[17]
The
Sonar reflectivity
The swim bladder of a fish can strongly reflect sound of an appropriate frequency. Strong reflection happens if the frequency is tuned to the volume resonance of the swim bladder. This can be calculated by knowing a number of properties of the fish, notably the volume of the swim bladder, although the well-accepted method for doing so[19] requires correction factors for gas-bearing zooplankton where the radius of the swim bladder is less than about 5 cm.[20] This is important, since sonar scattering is used to estimate the biomass of commercially- and environmentally-important fish species.
Deep scattering layer
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.[21]
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.[22][23] 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.[24] 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.[25] 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 tonnes, several times the annual 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.[26]
Human uses
In the East Asian culinary sphere, the swim bladders of certain large fishes are considered a food delicacy. In Chinese cuisine, they are known as fish maw, 花膠/鱼鳔,[27] and are served in soups or stews.
The vanity price of a vanishing kind of maw is behind the imminent extinction of the
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.[30] In earlier times, they were used to make condoms.[31]
Swim bladder disease
Risk of injury
Many
Similar structures in other organisms
Gallery
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Swim bladder display in a Malacca shopping mall
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Fish maw soup
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Swim bladder disease has resulted in this female ryukin goldfish floating upside down
References
- ^ "More on Morphology". www.ucmp.berkeley.edu.
- ^ "Fish". Microsoft Encarta Encyclopedia Deluxe 1999. Microsoft. 1999.
- ^ a b Darwin, Charles (1859) Origin of Species Page 190, reprinted 1872 by D. Appleton.
- JSTOR 1445488.
- S2CID 11198182.
- ^ "Secretion Of Nitrogen Into The Swimbladder Of Fish. Ii. Molecular Mechanism. Secretion Of Noble Gases". Biolbull.org. 1981-12-01. Retrieved 2013-06-24.
- ISBN 9780073524238.
- PMID 21532967.
- S2CID 7395840.
- ISBN 9780415375627.
- PMID 20190038.
- ^ Encyclopedia of Evolution
- ^ Does the bowfin gas bladder represent an intermediate stage during the lung-to-gas bladder evolutionary transition?
- ^ Tracing the genetic footprints of vertebrate landing in non-teleost ray-finned fishes
- ^ Allometric growth in the extant coelacanth lung during ontogenetic development
- ^ S2CID 87285937.
- ^ Biology of Fishes
- ISBN 0-697-28654-1
- doi:10.1121/1.382009.
- PMID 23297876.
- ^ Ryan P "Deep-sea creatures: The mesopelagic zone" Te Ara - the Encyclopedia of New Zealand. Updated 21 September 2007.
- ISBN 9780131008472.
- ISBN 9780203885222.
- PMID 1251208.
- ISBN 978-0-12-547665-2.
- ^ R. Cornejo; R. Koppelmann & T. Sutton. "Deep-sea fish diversity and ecology in the benthic boundary layer". Archived from the original on 2013-06-01. Retrieved 2015-03-26.
- ISBN 9781556437656.
- . Retrieved 14 October 2022.
- ^ "'Extinction Is Imminent': New report from Vaquita Recovery Team (CIRVA) is released". IUCN SSC - Cetacean Specialist Group. 2016-06-06. Archived from the original on 2019-01-03. Retrieved 2017-01-25.
- ^ Bridge, T. W. (1905) [1] "The Natural History of Isinglass"
- PMC 1974678.
- ISBN 0-8348-0448-4
- ^ PMID 23055066.
- PMID 22745695.
- ISBN 9781441973115.
- S2CID 2687386.
t
Further references
- Bond, Carl E. (1996) Biology of Fishes, 2nd ed., Saunders, pp. 283–290.
- Pelster, Bernd (1997) "Buoyancy at depth" In: WS Hoar, DJ Randall and AP Farrell (Eds) Deep-Sea Fishes, pages 195–237, Academic Press. ISBN 9780080585406.