gill

The red gills of this common carp are visible as a result of a gill flap birth defect.

A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist. The microscopic structure of a gill presents a large surface area to the external environment.

Many microscopic aquatic animals, and some that are larger but inactive, can absorb adequate oxygen through the entire surface of their bodies, and so can respire adequately without a gill. However, more complex or more active aquatic organisms usually require a gill or gills.^{[citation needed]}

Gills usually consist of thin filaments of tissue, branches, or slender tufted processes that have a highly folded surface to increase surface area. A high surface area is crucial to the gas exchange of aquatic organisms as water contains only a small fraction of the dissolved oxygen that air does. A cubic meter of air contains about 250 grams of oxygen at STP. For a typical freshwater oxygen concentration of 5 parts per millon by mass, a cubic meter of water will contain 5 grams of oxygen. This is about 1/50th of the oxygen of the same volume of air.

With the exception of some aquatic insects, the filaments and lamellae (folds) contain blood or coelomic fluid, from which gases are exchanged through the thin walls. The blood carries oxygen to other parts of the body. Carbon dioxide passes from the blood through the thin gill tissue into the water. Gills or gill-like organs, located in different parts of the body, are found in various groups of aquatic animals, including mollusks, crustaceans, insects, fish, and amphibians.

Vertebrate gills

Freshwater Fish Gills magnified 400 times

The gills of vertebrates typically develop in the walls of the pharynx, along a series of gill slits opening to the exterior. Most species employ a countercurrent exchange system to enhance the diffusion of substances in and out of the gill, with blood and water flowing in opposite directions to each other. The gills are composed of comb-like filaments, the gill lamellae, which help increase their surface area for oxygen exchange.^[1]

When a fish breathes, it draws in a mouthful of water at regular intervals. Then it draws the sides of its throat together, forcing the water through the gill openings, so that it passes over the gills to the outside. Fish gill slits may be the evolutionary ancestors of the tonsils, thymus gland, and Eustachian tubes, as well as many other structures derived from the embryonic branchial pouches.^{[citation needed]}

Cartilaginous fish

Sharks and rays typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six or seven pairs. Adjacent slits are separated by a cartilaginous gill arch from which projects a long sheet-like septum, partly supported by a further piece of cartilage called the gill ray. The individual lamellae of the gills lie on either side of the septum. The base of the arch may also support gill rakers, small projecting elements that help to filter food from the water.^[2]

A smaller opening, the spiracle, lies in front of the first gill slit. This bears a small pseudobranch that resembles a gill in structure, but only receives blood already oxygenated by the true gills.^[2] The spiracle is thought to be homologous to the ear opening in higher vertebrates.^[3]

Most sharks rely on ram ventilation, forcing water into the mouth and over the gills by rapidly swimming forward. In slow-moving or bottom dwelling species, especially among skates and rays, the spiracle may be enlarged, and the fish breathes by sucking water through this opening, instead of through the mouth.^[2]

Chimaeras differ from other cartilagenous fish, having lost both the spiracle and the fifth gill slit. The remaining slits are covered by an operculum, developed from the septum of the gill arch in front of the first gill.^[2]

Bony fish

The red gills inside a detached tuna head (viewed from behind)

In bony fish, the gills lie in a branchial chamber covered by a bony operculum. The great majority of bony fish species have five pairs of gills, although a few have lost some over the course of evolution. The operculum can be important in adjusting the pressure of water inside of the pharynx to allow proper ventilation of the gills, so that bony fish do not have to rely on ram ventilation (and hence near constant motion) to breathe. Valves inside the mouth keep the water from escaping.^[2]

The gill arches of bony fish typically have no septum, so that the gills alone project from the arch, supported by individual gill rays. Some species retain gill rakers. Though all but the most primitive bony fish lack a spiracle, the pseudobranch associated with it often remains, being located at the base of the operculum. This is, however, often greatly reduced, consisting of a small mass of cells without any remaining gill-like structure.^[2]

Marine teleosts also use gills to excrete electrolytes. The gills' large surface area tends to create a problem for fish that seek to regulate the osmolarity of their internal fluids. Saltwater is less dilute than these internal fluids, so saltwater fish lose large quantities of water osmotically through their gills. To regain the water, they drink large amounts of seawater and excrete the salt. Freshwater is more dilute than the internal fluids of fish, however, so freshwater fish gain water osmotically through their gills.^[2]

Other vertebrates

An Alpine newt larva showing the external gills, which flare just behind the head

Lampreys and hagfish do not have gill slits as such. Instead, the gills are contained in spherical pouches, with a circular opening to the outside. Like the gill slits of higher fish, each pouch contains two gills. In some cases, the openings may be fused together, effectively forming an operculum. Lampreys have seven pairs of pouches, while hagfishes may have six to fourteen, depending on the species. In the hagfish, the pouches connect with the pharynx internally. In adult lampreys, a separate respiratory tube develops beneath the pharynx proper, separating food and water from respiration by closing a valve at its anterior end.^[2]

Tadpoles of amphibians have from three to five gill slits that don't contain actual gills. There is usually no spiracle or true operculum, though many species have an operculum-like structure. Instead of internal gills, they develop three feathery external gills that grow from the outer surface of the gill arches. Sometimes adults retain these, but they usually disappear at metamorphosis. Lungfish larvae also have external gills, as does the primitive ray-finned fish Polypterus, though the latter has a structure different than amphibians.^[2]

Branchia

Branchia (pl. branchiæ) is the Ancient Greek naturalists' name for gills. Galen observed that fish had multitudes of openings (foramina), big enough to admit gases, but too fine to give passage to water. Pliny the Elder held that fish respired by their gills, but observed that Aristotle was of another opinion.^[4] The word branchia comes from the Greek βράγχια, "gills", plural of βράγχιον (in singular, meaning a fin).^[5]

Invertebrate gills

A live individual of the sea slug Pleurobranchaea meckelii. The gill (or ctenidium) is visible in this view of the right-hand side of the animal

Respiration in the Echinodermata (includes starfish and sea urchins) is carried out using a very primitive version of gills called papulae. These thin protuberances on the surface of the body contain diverticula of the water vascular system. Crustaceans, molluscs, and some insects have gills that are tufted or plate-like structures at the surface of the body.

Caribbean hermit crabs have modified gills that allow them to live in humid conditions

The gills of other insects are tracheal, and also include both thin plates and tufted structures, and, in the larval dragon fly, the wall of the caudal end of the alimentary tract (rectum) is richly supplied with tracheae as a rectal gill. Water pumped into and out of the rectum provides oxygen to the closed tracheae. Aquatic insects use a tracheal gill, which contains air tubes. The oxygen in these tubes is renewed through the gills.

Physical gills

Physical gills are a type of structural adaptation common among some types of aquatic insects, which holds atmospheric oxygen in an area with small openings called spiracles. The structure (often called a plastron) typically consists of dense patches of hydrophobic setae on the body, which prevent water entry into the spiracles. The physical properties of the interface between the trapped air bubble and surrounding water accomplish gas exchange through the spiracles, almost as if the insect were in atmospheric air. Carbon dioxide diffuses into the surrounding water due to its high solubility, while oxygen diffuses into bubbles as the concentration within the bubble has been reduced by respiration, and nitrogen also diffuses out as its tension has been increased. Oxygen diffuses into the bubble at a higher rate than Nitrogen diffuses out. However, water surrounding the insect can become oxygen-depleted if there is no water movement, so many aquatic insects in still water actively direct a flow of water over their bodies.

The physical gill mechanism allows aquatic insects with plastrons to remain constantly submerged. Examples include many beetles in the family Elmidae, aquatic weevils, and true bugs in the family Aphelocheiridae.^{[citation needed]}

See also

• Aquatic respiration
• Book lung
• Gill raker
• Gill slit
• Lung
• Artificial gills (human)

References

This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (May 2011)

External links

• Fish Dissection - Gills exposed Australian Museum. Updated: 11 June 2010. Retrieved 16 January 2012.

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