Swim Bladders and the Origin of Lungs

Seth Andrew J. Salih

Swim Bladders and Lungs

Every osteichthyan from fish to humans develops an unpaired evagination from the foregut that becomes pneumatic sacs The pneumatic sacs become either swim bladders or lungs These sacs fill with gases derived directly or indirectly from the environment

Swim Bladders and Lungs

Swim Bladders and Lungs

Pneumatic sacs do not imply a function Swim bladders and lungs suggest functions for buoyancy and respiration Air sacs of fishes called swim bladder and affect buoyancy Air sacs of tetrapods referred to as lungs

Swim Bladders and Lungs

A clade of tailed amphibians Bottom dwellers

Latimeria
Without pneumatic (air) sacs

Marine Teleosts

Swim Bladders
After the budlike anlage (foundation) for a swim bladder evaginates, the resulting duct may retain the connection to the foregut or may close during later development

Fishes with ducts that remain open are physostomous Examples are: Chondrosteans,basal neopterygians, the 3 living dipnoans and some teleosts

Swim Bladders
Fishes with ducts that close are physoclistous Most teleosts are physoclistous.

Swim Bladders
Physoclistous Physostomous

Swim Bladders
Swim Bladders may be paired or unpaired. In chondrosteans and dipnoans, the unpaired duct leads from the ventral aspect of the esophagus In neopterygians, it has shifted toward the dorsal side Though less frequently, the duct leads from the pharynx or stomach

Swim Bladders
Swim bladders lie close to kidneys and are retroperitoneal in position As the embryonic anlagen grow longer, they push their way caudad in the roof of the coelom between the embryonic parietal peritoneum and body wall However, adult swim bladders may bulge into the roof of the coelom

Swim Bladders

Swim Bladders
Swim bladders serve as hydrostatic organs, respiratory organs, organs participating in sound detection, and organs of communication Their capacity for gases can be increased/decreased due to the elastic and smooth muscles in the bladder walls

Swim Bladders
In teleosts, they function mainly as hydrostatic organs that maintain an appropriate depth or hover over a specific location To do this, a fish must attain a body density equal to that of the displaced water at that depth This is done by controlling the volume of gas in the bladder. A difference in density will affect the buoyancy, causing the fish to rise/sink

Swim Bladders

Swim Bladders
However, a fish must use its fins to compensate for the disturbances in the water Some fishes make daily changes in depth and body density as a correlate of the tome of day The gas in the bladder comes from the blood and is actively transported to the lumen of the bladder from the red gland, a localized rete (network) of small arteries in the bladder lining

Swim Bladders
The gas is eventually resorbed into the bloodstream of the bladder near its caudal end in a pocketlike area of modified epithelium The pocket can be closed off from the main cavity by a sphincter during passage of gas into the lumen of the bladder and relaxed when resorption of gas is taking place In physostomes, the gas may be bubbled to the exterior through the mouth

Swim Bladders
The gases in the bladder differ among fishes Some contain 99% nitrogen Some, up to 87% oxygen All contain traces of carbon dioxide and argon

In deepwater fish, nitrogen may be transported from the blood into the lumen of the bladder

Swim Bladders
In physostomous fishes, swim bladders function as lungs to one degree Air is gulped at the surface of the water and an oropharyngeal pump forces it into the swim bladder Air depleted of oxygen is expelled by the vacuum created by lowering the floor of the oropharyngeal cavity Expulsion is facilitated by the elasticity of the bladder wall, its smooth muscle musculature and the pressure of the surrounding water Air is then bubbled through the mouth into the water

Swim Bladders
2 of 3 genera of lungfishes, Propterus and Lepidosiren, require continual gulping of air for survival because their gills are incapable of producing sufficient O2 They drown if held underwater for a certain length of time Their swim bladders act as lungs during tropical summers, when swamps dry up, and allow them to survive Additionally, these fish also absorb O2 from the skin

Swim Bladders
The swim bladder of the other dipnoan (Neoceratodus), 2 genera of ray-finned African freshwater fishes, Polypterus, Calamoichthys, and relict basal neopterygians (Amia and gars) function as a lung only when the O2 content of water is so low that gills cannot produce the respiratory quota of the organism

Swim Bladders
Polypterus and Calamoichthys True lungfishes
Smooth lining Supplied by arteries from the 6th embryonic aortic arch (along with Amia) Lined with low septa and exhibit thousands of air sacs Supplied by arteries from the 6th embryonic aortic arch (along with Amia)

Swim Bladders
Like tetrapods, the venous return in dipnoans is directly to the left atrium of the heart Some teleosts use their swim bladders for sound detection In Cypriniformes, small bones, weberian ossicles, connect the anterior end of the bladder with the sinus impar, an extension of the perilymphatic space of the inner ear. In Clupeiformes, a thin-walled anterior extension of the bladder makes direct contact with the inner ear.

Swim Bladders
In some fishes, contraction of striated muscles attached to the bladder causes it to emit thumping sounds, or forces air back and forth between chambers separated by muscular sphincters within the bladder This produces croaking or grunting sounds for communication

Swim Bladders
Bottom feeders have degenerate swim bladders Eliminating this hydrostatic organ optimized their body density, enabling them to hover close to food with little energy expenditure An analogous condition is in salamanders who live in swift mountain streams and whose lungs are only a few mm or absent Optimal body density and the saving of energy are the benefits

Swim Bladders
However, no member of the plethodont family has lungs. These urodeles breath through their skin

Origin of Lungs
The similarities between swim bladders and lungs suggest that they may be the same organs During the Devonian, the swamps were warm ad stagnant, causing them to be low in O2. This condition may have caused the development of aerial respiration among organisms At the time, most rhipidistians had pneumatic sacs

Origin of Lungs
A pneumatic sac was functioning in aerial respiration before craniates ventured onto land The closure of the pneumatic duct in physoclistous fishes is probably derived from a more primitive open-duct condition