- Key People:
- Sir John Graham Kerr
There are a number of fishes that, in addition to or in place of gill breathing, have developed special organs through which they can breathe atmospheric air at the water surface. This occurs almost exclusively in freshwater fishes. In lungfishes these organs are, both in function and in structure, primitive lungs like those of amphibians. The name lungfish is thus well applied: these fishes have lungs that are derived from the swim bladder (an organ used for buoyancy in most bony fishes), which is connected to the alimentary tract. The inner surfaces of these air-breathing organs are covered with a great number of honeycomb-like cavities supplied with fine blood vessels. As in terrestrial higher vertebrates, gas exchange takes place in tiny air vesicles. Also as in terrestrial vertebrates, there is a separate pulmonary circulation.
In order to breathe, the fish swims upward and positions its head so that the tip of the snout barely touches the water surface. The mouth is then opened wide, and the fish sucks in air from just above the water—a process often accompanied by a characteristic sound. The Australian lungfish reportedly breathes air through the nasal openings, the mouth remaining closed. In contrast to the more advanced bony fishes, lungfishes have a particular opening (choana) that connects the nasal cavity with the mouth.
In the Australian lungfish, gill breathing predominates at least some of the time—namely, in times of normal water level when the water is well oxygenated. At such times the fish rises less often to the surface to breathe atmospheric air. When the water level goes down, which usually occurs in August or September, the fish is often found in isolated waterholes in which the oxygen content is greatly reduced. Other fishes in such pools often die from lack of oxygen, but the lungfish survives, having changed over to the breathing of atmospheric air. During such a dry period the Australian lungfish surfaces about every 40 to 50 minutes for air. African lungfishes surface for air about every 30 minutes or, in some cases, at longer intervals.
Physiology and biochemistry
African lungfishes burrow into the bottom of a riverbed or lake bed for their “dry sleep,” or estivation (see dormancy). After burying themselves, they become encased in a mucous sheath that gradually hardens. Here they spend the dry season, during which the waterline becomes lower and the riverbed or lake bed finally dries out. The African lungfish generally digs in and encysts in this manner, even if there is sufficient time to swim to deeper waters. African lungfishes also burrow into mud and ensheathe themselves under experimental conditions. They have been kept alive in such an induced state for more than two years. The South American lungfish also burrows into the mud in times of water shortage, but it forms no protective sheath. However, the Australian lungfish never buries itself in this manner. During prolonged estivation, African lungfishes may accumulate high concentrations of urea in the body.
Studies have shown that a substance that inhibits the fish’s normal metabolism induces the dry sleep of the African lungfish. Extracts from the brains of such sleeping fish injected into rats have caused them to become lethargic; in addition, the body temperature of the rats falls 5 °C (9 °F), and the metabolic rate falls 33 percent. The day after receiving such injections, the rats stop eating. It is believed that the material responsible for this effect is a proteinlike substance.
Evolution
The oldest Dipnoi, from the Early Devonian, possessed skull and dental features that were characteristically dipnoid but also had many features in common with another sarcopterygian group called the crossopterygians, such as the coelacanths. The Dipnoi were abundant until Triassic times (about 251.9 million to 201.3 million years ago), after which their numbers decreased.
Dipterus, one of the oldest lungfish, had leaflike pectoral and pelvic fins similar to those of the modern Australian lungfish, and it seems reasonable to assume that early forms also had functional lungs comparable to those of species living today. Hardened sections of clay, cylindrical in shape, have been found in deposits dating to Pennsylvanian and Permian times (about 323.2 million to 251.9 million years ago). Remains of the dipnoid Gnathorhiza, closely allied to the extant African and South American species, were embedded in the clay. Their discovery in such a setting strongly suggests that these dipnoids passed unfavourable conditions buried in mud.
An evolutionary line can be traced from Dipterus to Neoceratodus, the extant Australian genus. Scaumenacia and Phaneropleuron, common forms in the Late Devonian (about 382.7 million to 358.9 million years ago), exhibited a much-reduced first dorsal fin (the first fin forward on the back); the second dorsal fin was enlarged and had shifted further toward the tail. Lungfish of Permian times showed an apparent fusion of the fins along the back and the rest of the vertical midline. This extended fin tapered to a point at the tip of the tail and also occurs in modern lungfishes. Various side branches also emerged in the evolution of the Dipnoi; however, none of these have survived to modern times.
Some scientists claim that walking and bounding behaviours common to tetrapods (limbed vertebrates and their descendants) evolved first in sarcopterygian fishes. They cite as evidence the ability of a modern African lungfish, Protopterus annectens, to propel itself along a substrate, such as the bottom of a pool, using its long fins for support. The fins contain no digits, yet the animal may lift its body up from the substrate by bending them to function like footlike appendages. The animal routinely employs a quadrupedal gait; however, it can also adopt a bipedal gait at times. Since tetrapods arose from sarcopterygian fishes, the presence of walking and bounding behaviours in lungfishes may be proof that these traits evolved before the emergence of tetrapods.