Structure and Function
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 pressure 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 and filling the swim bladder. 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, fish have to rise to the surface to fill up their swim bladders, however, in later stages the connection 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. 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.
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.
In some fish, mainly freshwater species (e.g. common carp, wels catfish), the swim bladder is connected to the labyrinth of the inner ear by the Weberian apparatus, a bony structure derived from the vertebrae, which provides a precise sense of water pressure (and thus depth), and improves hearing.
Read more about this topic: Swim Bladder
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