2.3.

Plaukiojamoji pūslė
Kaip ir daugelio kitų žuvų, virš ešerio žarnyno yra tam tikra plonasienė išauga, pripildyta ddujų mišinio, –
plaukiojamoji pūslė. Suaugusio ešerio pūslė su žarnynu nesijungi, bet jo lervos ir kai kurių kitų žuvų
(pavyzdžiui, kuojos arba karpio) tarp žarnyno ir pūslės visą amžių išlieka nedidelis latakėlis.

Plaukiojimo pūslė – daugelio kaulinių žuvų hidrostatinis organas, leidžiantis žuviai valdyti kūno plūduriavimą
keičiant joje esančių dujų kiekį
. Kai pūslė sumažėja žuvis pasunkėja ir grimzta gilyn. Tai padeda greitai pasinerti.Atitinkamai ,kai pūslė
padidėja žuvis truputį palengvėja ir kyla aukštyn. Kol žuvis būna tame pačiame gylyje, pūslės apimtis
nesikeičia.

Swim bladder
fish anatomy
WRITTEN BY: The Editors of Encyclopaedia Britannica
Alternative Title: air bladder
Swim bladder, also called air bladder, buoyancy organ possessed by most bony fish. The swim bladder is located in the body
cavity and is derived from an outpocketing of the digestive tube. It contains gas (usually oxygen) and functions as a hydrostatic,
or ballast, organ, enabling the fish to maintain its depth without floating upward or sinking. It also serves as a resonating
chamber to produce or receive sound. In some species the swim bladder contains oil instead of gas. In certain primitive fish it
functions as a lung or respiratory aid instead of a hydrostatic organ. The swim bladder is missing in some bottom-dwelling and
deep-sea bony fish (teleosts) and in all cartilaginous fish (sharks, skates, and rays).

Teleost fish in cross section.Encyclopædia Britannica, Inc.

Swim bladder
From Wikipedia, the free encyclopedia
For Fish bladder symbol, see fish bladder.

The swim bladder of a rudd

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.
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, but scientists now believe that the swim bladder derived from a more primitive lung.
[citation needed]

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

Swim bladder from a bony (teleost) fish

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.
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]
Evolution[edit]

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.[10] 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.[10] 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.[11]
Deep scattering layer[edit]

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.[12]
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.[13][14] 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.[15] 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.
[16]
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.[17]
Human uses[edit]
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,[18] 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 breed. Only found in Mexico's Gulf of California, the once numerous vaquita now number less
than 60 in total. Vaquita die in gillnets[19] 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. In earlier times they were used to
make condoms.[20]
Swim bladder disease[edit]
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.[21]
Risk of injury[edit]
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.[22] 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.[22]
Investigators used the High-Intensity-Controlled Impedance Fluid Filled (HICI-FT), a stainless-steel wave tube
with a electromagnetic shaker. It simulates high-energy sound waves in aquatic far-field, plane-wave acoustic
conditions.[23][24]
Similar structures in other organisms[edit]
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.[25]

https://www.youtube.com/watch?v=6q7NQ7XrzS8

https://www.youtube.com/watch?v=TJN3gJoZqlY

https://www.youtube.com/watch?v=B-T4ORXLgLI

https://www.youtube.com/watch?v=zRa5M2Lmqak

https://www.youtube.com/watch?v=3DyRTDDurTM