Swim 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) to control their buoyancy, and thus to stay at their current water depth without having to waste energy in swimming. 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. Charles Darwin, 1859Swim bladder display in a Malacca shopping mallFish maw soupSwim bladder disease has resulted in this female ryukin goldfish floating upside down 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) to control their buoyancy, and thus to stay at their current water depth without having to waste energy in swimming. 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. Darwin reasoned that the lung in air-breathing vertebrates had derived from a more primitive swim bladder. In the embryonic stages, some species, such as redlip blenny, 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. 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. 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 in the swim bladder which accounts for the high pressures of other gases as well. 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.