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Gills and lungs are the two structures commonly used by animals for respiration. Both are characterized by large amounts of surface area that function in gas exchange. The difference between them is that gills involve external extensions from the body surface, whereas lungs possess internal foldings. Gills have evolved independently several times in a variety of animal groups.

Among the annelids, certain species of terrestrial worms have long slender, branching gills which extend from the body. Horseshoe crabs possess structures known as book gills, which are actually modified appendages that function in gas exchange. Crustaceans also have gills that have been modified from thoracic or abdominal appendages.

Gill structures are highly diverse among crustaceans. Generally, the more aquatic crustaceans have more elaborate gills, while the more terrestrial species are characterized by simplified gills. That is a consequence of the greater availability of oxygen on land, from the air, than in water.

In echinoderms, the group that includes starfish, the large surface area provided by the many appendages and by the tube feet are used in gas exchange. However, some species supplement these with gills around the oral cavity. Mollusks possess gills within their mantles. These are oriented to face the water current. Vertebrates such as salamanders are also characterized by external gills, which in their case are filamentous structures that extend from the head region. In some species, only the aquatic larvae have gills. However, in many species that remain aquatic their entire lives, the gills may be retained into adulthood.

Fish also use gills in gas exchange. The gills of fish are supported by a series of bony gill arches. The gill arches lie between the gill clefts, through which oxygenated water flows. The gill arches support tissue that includes the tiny blood vessels which carry in deoxygenated blood and carry away oxygenated blood, as well as the gill filaments, where gas exchange actually occurs. The gill filaments each have numerous secondary gill lamellae that further increase the surface area available for gas exchange.

Fish that have high energy demands, such as those which swim quickly, have more surface area associated with their gills. The operculum of bony fishes is a covering that protects the entire gill area. It also covers the operculum chamber, which is essential to the process of pumping water over the gills.

Gas exchange across the gills occurs in what is described as a counter current fashion. This is a very efficient mode of gas exchange because water flows in one direction and blood flows in the other. The consequence of countercurrent flow is that well-oxygenated blood encounters well-oxygenated water, and is able to pull more oxygen from the water, while less-oxygenated blood encounters less-oxygenated water. Oxygenation thus occurs along the entire pathway where the water and blood are juxtaposed.

Fish push oxygen-rich water across their gills by one of two methods. In ram ventilation, fast-swimming species such as sharks swim with their mouths open. Water is forced into the mouth and out over the gills.

Most bony fish species, however, employ a second method, buccal and opercular pumping. In this method, serial expansions and contractions of the mouth cavity and the opercular cavity occur, resulting in the continuous flow of water over the gills. First, the mouth, or buccal, cavity expands, drawing water in. Then, the mouth closes and the buccal cavity contracts. This forces water to flow over the gills into the opercular cavity. At the same time, the opercular cavity expands, which draws more water in from the buccal cavity. Water exits through the operculum.

Some fish will use buccal and opercular pumping while swimming slowly and ram ventilation when swimming faster. Ram ventilation is essential in some species, which suffocate if they are not able to swim fast enough.

The gill arches have played a crucial role in vertebrate evolution. Over evolutionary time, they have been modified to form other essential skeletal structures. The vertebrate jaw, which characterizes all vertebrates aside from lamprey and hagfish, was modified from a single pair of gill arches.

In mammals there has been a further modification of this pair of gill arches. Mammals have evolved a "new jaw," and the original bones of the jaw joint now function as middle-ear bones in mammals. The incus and malleus are both homologous to the jaw bones in other vertebrates, which are homologous to gill arches in primitive fish groups. The middle-ear bones are involved in conducting sound between the eardrum and the inner ear, where neural processing occurs. The third mammalian middle-ear bone, the stapes, occurs in all terrestrial vertebrates, and is also derived from the gill arches.

see also Blood; Respiratory System.

Jennifer Yeh


Gould, James L., and William T. Keeton. Biological Science, 6th ed. New York: W. W. Norton, 1996.

Hildebrand, Milton, and Viola Hildebrand. Analysis of Vertebrate Structure. New York: John Wiley, 1994.

Withers, Philip C. Comparative Animal Physiology. Fort Worth, TX: Saunders College Pub., 1992.

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gill1 / gil/ • n. (often gills) 1. the paired respiratory organ of fishes and some amphibians, by which oxygen is extracted from water flowing over surfaces within or attached to the walls of the pharynx. ∎  an organ of similar function in an invertebrate animal. 2. the vertical plates arranged radially on the underside of mushrooms and many toadstools. 3. the wattles or dewlap of a fowl. ∎  (gills) the flesh below a person's jaws and ears: we stuffed ourselves to the gills with scrambled eggs and toast. • v. [tr.] 1. gut or clean (a fish). 2. catch (a fish) in a gill net. PHRASES: green around (or at) the gills (of a person) sickly-looking.DERIVATIVES: gilled adj. [in comb.] a six-gilled shark. gill2 / jil/ • n. a unit of liquid measure, equal to a quarter of a pint. gill3 / gil/ • n. Brit. a deep ravine, esp. a wooded one. ∎  a narrow mountain stream. gill4 / jil/ (also jill) • n. 1. archaic a young woman; a sweetheart. 2. a female ferret. Compare with hob2 (sense 1).

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gills, external respiratory organs of most aquatic animals. In fishes the gills are located in gill chambers at the rear of the mouth (pharynx). Water is taken in through the mouth, is forced through openings called gill slits, and then passes through the gill clefts, spaces between the ranks of delicate gills, bathing them continuously. Each gill is composed of numerous threadlike gill filaments containing capillaries enclosed in a thin membrane; oxygen is absorbed from the passing water and carbon dioxide is discharged. The gills, which may be platelike or tufted, are attached to the outer edges of a series of paired cartilaginous or bony gill (or branchial) arches. Gill rakers, bony comblike projections on the inner edge of the arches, strain solid material from the water, preventing it from passing out through the gill slits and directing it down the esophagus. Gill rakers are present in all fishes except those that feed on large organisms. In primitive fishes (e.g., the shark) the gill slits are exposed; in the bony fishes they are protected by an operculum, or gill cover. In the higher aquatic invertebrates the gills protrude from the body surface and contain extensions of the vascular system. In the crustaceans these external gills are covered by a protective carapace, part of the shell; in the echinoderms they are branched appendages extending from various parts of the body. In the mollusks the gills (called ctenidia) are internal and are located inside the mantle cavity. Horseshoe crabs have gill books, which are membranous flaps like the pages of a book. Amphibians breathe by means of external gills in their aquatic larval stage; a few forms retain the gills after metamorphosing into terrestrial adults. Aquatic insect larvae accomplish the oxygen–carbon-dioxide exchange by means of tracheal gills, projections from the walls of the air tubes (tracheae); these gills disappear when the insect leaves the water. The embryos of all higher vertebrates pass through a stage in which rudimentary gill slits occur, but these never become functional and disappear as the embryo continues to develop.

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Gillbill, Brazil, brill, Camille, chill, cookchill, dill, distil (US distill), downhill, drill, Edgehill, Estoril, fill, freewill, frill, fulfil (US fulfill), Gill, goodwill, grill, grille, hill, ill, instil, kill, krill, mil, mill, nil, Phil, pill, quadrille, quill, rill, Seville, shill, shrill, sill, skill, spadille, spill, squill, still, stock-still, swill, thill, thrill, till, trill, twill, until, uphill, will •hwyl • bank bill • handbill • waxbill •playbill, waybill •cranesbill • sibyl • crossbill • sawbill •hornbill • storksbill • shoebill •spoonbill • duckbill • razorbill •gerbil • wind chill • Churchill • idyll •daffodil • back-fill • landfill • monofil •fibrefill (US fiberfill) • chlorophyll •bluegill

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1. (in zoology) A respiratory organ used by aquatic animals to obtain oxygen from the surrounding water. A gill consists essentially of a membrane or outgrowth from the body, with a large surface area and a plentiful blood supply, through which diffusion of oxygen and carbon dioxide between the water and blood occurs. Fishes have internal gills, formed as outgrowths from the pharynx wall and contained within gill slits. Water entering the mouth is pumped out through these slits and over the gills. The gills of most aquatic invertebrates and amphibian larvae are external gills, which project from the body so that water passes over them as the animal moves.

2. (in botany) One of the ridges of tissue that radiate from the centre of the underside of the cap of mushrooms. The spores are produced on these gills.

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gill The respiratory organ of an aquatic animal, often complex in form, consisting of an outgrowth from the body surface or an internal layer of modified gut, past which water flows. It provides a large surface area, well supplied with blood vessels, across whose walls oxygen and carbon dioxide diffuse. It is the chief organ of respiration in fish, in which the gills usually are supported by four or more pairs of bony or cartilaginous gill arches. In sharks each gill is more or less surrounded by a separate gill chamber, but in bony fish the gills share a common gill (branchial) chamber, one on each side of the head. Each gill arch bears rows of plate-like filaments, in turn carrying secondary lamellae, which are capable of absorbing oxygen from the water flowing over the surface.

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gill A membranaceous, blade-like structure (lamella) which bears the hymenium in the fruit body of a mushroom or other agaric. Gills are usually arranged radially on the underside of the pileus.

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gill Organ through which most fish, some larval amphibians and many aquatic invertebrates obtain oxygen from water. When a fish breathes, it opens its mouth, draws in water and shuts its mouth again. Water is forced through the gill slits, over the gills, and out into the surrounding water. Oxygen is absorbed into small blood vessels, and at the same time, waste carbon dioxide carried by the blood diffuses into the water.

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gill2 rocky cleft, ravine XI (in place-names), XIV (in literature); narrow stream XVII. — ON. gil deep glen. The fanciful sp. ghyll was introduced by Wordsworth.