A gill ( ) is a specialized respiratory organ that many aquatic animals use for aquatic gas exchange, i.e. to extract dissolved oxygen from water and to excrete carbon dioxide. Branchia (pl.: branchiae, from Ancient Greek βράγχια) is the zoologists' academic name for gills. The gills of some semi-aquatic clades (e.g. crabs, terrestrial hermit crabs and amphibious fishes), have also adapted to allow air respiration (breathing) on land provided the gills are kept moist. In some terrestrial chelicerates species (tetrapulmonates such as spiders and scorpions), their ancestral gills have evolved into fully air-breathing book lungs. In some planktivorous bony fish species (e.g. silver and bighead carps), the gills are also used as a filter feeding organ via comb-like projections called gill rakers. The microscopic structure of a gill presents a large surface area in contact with the external environment, which allows optimal diffusion. With the exception of some aquatic insects, the gills of aquatic animals have filaments and lamellae (folds) that contain blood vessels or coelomic fluid, from which gases are exchanged into and out of blood/hemolymph through the thin gill walls before being distributed by the circulatory system to other parts of the body. Gills or gill-like organs, located in different parts of the body, are found in various groups of aquatic animals, including molluscs, crustaceans, xiphosurans, aquatic insects, polychaetes and most aquatic vertebrates (fish and amphibian tadpoles). Semi-terrestrial marine animals such as crabs and mudskippers have gill chambers in which they store water, enabling them to temporarily survive on the reservoir of dissolved oxygen when they are on land.
History 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. The word branchia comes from the Greek βράγχια, "gills", plural of βράγχιον (in singular, meaning a fin).
Function Many microscopic aquatic animals, and some larger but inactive ones, can absorb sufficient oxygen through the entire surface of their bodies, and so can respire adequately without gills. However, more complex or more active aquatic organisms usually require one gill or more. Many invertebrates, and even amphibians, use both the body surface and gills for gaseous exchange. Gills usually consist of thin filaments of tissue, lamellae (plates), branches, or slender, tufted processes that have a highly folded surface to increase surface area. The delicate nature of the gills is possible because the surrounding water provides support. The blood or other body fluid must be in intimate contact with the respiratory surface for ease of diffusion. A high surface area is crucial to the gas exchange of aquatic organisms, as water contains only a small fraction of dissolved oxygen compared to the oxygen content of air, and it diffuses more slowly. A cubic meter of air contains about 275 grams of oxygen at STP. Fresh water holds less than 1/25th the oxygen content of air, the dissolved oxygen content being approximately 8 cm3/L compared to the oxygen content of air which is 210 cm3/L. Water is 777 times more dense than air and is 100 times more viscous. Oxygen has a diffusion rate in air 10,000 times greater than in water. The use of sac-like lungs to remove oxygen from water would not be efficient enough to sustain life. Rather than using lungs, "[g]aseous exchange takes place across the surface of highly vascularised gills over which a one-way current of water is kept flowing by a specialised pumping mechanism. The density of the water prevents the gills from collapsing and lying on top of each other; [such collapse] happens when a fish is taken out of water." Usually water is moved across the gills in one direction by the current, by the motion of the animal through the water, by the beating of cilia or other appendages, or by means of a pumping mechanism. In fish and some molluscs, the efficiency of the gills is greatly enhanced by a countercurrent exchange mechanism in which the water passes over the gills in the opposite direction to the flow of blood through them. This mechanism is very efficient and as much as 90% of the dissolved oxygen in the water may be recovered.
Vertebrates
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. 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 it passes over the gills to the outside. Fish gill slits may be the evolutionary ancestors of the thymus glands, parathyroid glands, as well as many other structures derived from the embryonic branchial pouches.
Fish
The gills of fish form a number of slits connecting the pharynx to the outside of the animal on either side of the fish behind the head. Originally there were many slits, but during evolution, the number reduced, and modern fish mostly have five pairs, and never more than eight.
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