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By adjusting the volume of gas in the swim bladder, a fish can achieve neutral buoyancy and remain suspended indefinitely at any depth with no muscular effort. There are severe technical problems, however. If the fish descends to a greater depth, the swim bladder gas is compressed so that the fish becomes heavier and tends to sink. Gas must be added to the bladder to establish a new equilibrium buoyancy. If the fish swims up, the gas in the bladder expands, making the fish lighter. Unless gas is removed, the fish will rise with ever increasing speed while the bladder continues to expand, until it pops helplessly out of the water. There are two ways fish adjust gas volume in the swim bladder. The less specialized fishes (trout, for example) have a pneumatic duct that connects the swim bladder to the esophagus ; these forms must come to the surface and gulpair to charge the bladder and obviously are restricted to relatively shallow depths. More specialized teleost shave lost the pneumatic duct. Gas exchange depends on two highly specialized areas: a gas gland that secretes gas into the bladder and a resorptive area, or oval, that can remove gas from the bladder. The gas gland contains a remarkable network of blood vessels (rete mirabile 迷网) arranged so that a vast number of arteries and veins in a tight bundle run in opposite directions to each other. This is called countercurrent flow, and in some way, not yet understood, this arrangement makes possible a tremendous multiplication of gas concentration inside the swim bladder. The amazing effectiveness of this device is exemplified by a fish living at a depth of 8,000 feet. To keep the bladder inflated, the gas inside (mostly oxygen, but also variable amounts of nitrogen, carbon dioxide, carbon monoxide, and argon) must have a pressure exceeding 240 atmospheres. Yet the oxygen pressure in the fishs blood cannot exceed one-fifth atmosphere--equal to the oxygen pressure at the sea surface. Physiologists suggest tha
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