270:, the dissolved oxygen content is approximately 8 cm/L compared to that of air which is 210 cm/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, "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, which is what happens when a fish is taken out of water."
438:, 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 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.
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pairs of pouches, while hagfishes may have six to fourteen, depending on the species. In the hagfish, the pouches connect with the pharynx internally and a separate tube which has no respiratory tissue (the pharyngocutaneous duct) develops beneath the pharynx proper, expelling ingested debris by closing a valve at its anterior end.
727:, and nitrogen also diffuses out as its tension has been increased. Oxygen diffuses into the air film 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 such insects in still water actively direct a flow of water over their bodies.
714:
on the body, which prevent water entry into the spiracles, but may also involve scales or microscopic ridges projecting from the cuticle. The physical properties of the interface between the trapped air film and surrounding water allow gas exchange through the spiracles, almost as if the insect were
403:
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
480:
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
441:
The gill arches of bony fish typically have no septum, so 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 spiracles, the pseudobranch associated with them often remains, being located at the
219:
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 a gill or gills. Many invertebrates, and
529:
have from three to five gill slits that do not contain actual gills. Usually no spiracle or true operculum is present, though many species have operculum-like structures. Instead of internal gills, they develop three feathery external gills that grow from the outer surface of the gill arches.
341:
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.
273:
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
314:
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
588:, and some aquatic insects have tufted gills or plate-like structures on the surfaces of their bodies. Gills of various types and designs, simple or more elaborate, have evolved independently in the past, even among the same class of animals. The segments of
470:). Conversely, fresh water contains less osmolytes than the fish's internal fluids. Therefore, freshwater fishes must utilize their gill ionocytes to attain ions from their environment to maintain optimal blood osmolarity.
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of their internal fluids. Seawater contains more osmolytes than the fish's internal fluids, so marine fishes naturally lose water through their gills via osmosis. To regain the water, marine fishes drink large amounts of
278:
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.
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is a type of structural adaptation occurring among some aquatic arthropods (primarily insects), a form of inorganic gill which holds a thin film of atmospheric oxygen in an area with small openings called
239:. 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.
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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
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Aquatic arthropods usually have gills which are in most cases modified appendages. In some crustaceans these are exposed directly to the water, while in others, they are protected inside a
307:
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
675:, but the air tubes are sealed, commonly connected to thin external plates or tufted structures that allow diffusion. The oxygen in these tubes is renewed through the gills. In the
1148:
Laurin M. (1998): The importance of global parsimony and historical bias in understanding tetrapod evolution. Part I-systematics, middle ear evolution, and jaw suspension.
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many of which carry gills. Sponges lack specialised respiratory structures, and the whole of the animal acts as a gill as water is drawn through its spongy structure.
622:. A current of water is maintained through the gills for gas exchange, and food particles are filtered out at the same time. These may be trapped in
449:
also use their gills to excrete osmolytes (e.g. Na⁺, Cl). The gills' large surface area tends to create a problem for fish that seek to regulate the
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typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six pairs with the
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base of the operculum. This is, however, often greatly reduced, consisting of a small mass of cells without any remaining gill-like structure.
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690:) is richly supplied with tracheae as a rectal gill, and water pumped into and out of the rectum provides oxygen to the closed tracheae.
1167:"The Multifunctional Fish Gill: Dominant Site of Gas Exchange, Osmoregulation, Acid-Base Regulation, and Excretion of Nitrogenous Waste"
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differ from other cartilagenous fish, having lost both the spiracle and the fifth gill slit. The remaining slits are covered by an
1343:
1211:
Florian
Witzmann; Elizabeth Brainerd (2017). "Modeling the physiology of the aquatic temnospondyl Archegosaurus decheni from the
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that resembles a gill in structure, but only receives blood already oxygenated by the true gills. The spiracle is thought to be
102:, have adapted to allow respiration on land provided they are kept moist. The microscopic structure of a gill presents a large
730:
The inorganic gill mechanism allows aquatic arthropods with plastrons to remain constantly submerged. Examples include many
377:, projections into the pharyngeal cavity that help to prevent large pieces of debris from damaging the delicate gills.
1502:
1337:(1999). "Morphological structures and vertical distribution in the soil indicate facultative plastron respiration in
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1419:"The marine-associated lifestyle of ameronothroid mites (Acari, Oribatida) and its evolutionary origin: a review"
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have gill chambers in which they store water, enabling them to use the dissolved oxygen when they are on land.
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373:, which the individual lamellae of the gills lie on either side of. The base of the arch may also support
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534:. Examples of salamanders that retain their external gills upon reaching adulthood are the
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being the only cartilaginous fish exceeding this number. Adjacent slits are separated by a
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that connect to the tracheal system. The plastron typically consists of dense patches of
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even amphibians, use both the body surface and gills for gaseous exchange.
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of
Germany". Fossil Record. 20 (2): 105–127. doi:10.5194/fr-20-105-2017.
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Evans, David H.; Piermarini, Peter M.; Choe, Keith P. (January 2005).
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Still, some extinct tetrapod groups did retain true gills. A study on
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of aquatic organisms, as water contains only a small fraction of the
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1016:"Developmental and evolutionary origins of the pharyngeal apparatus"
1128:. Philadelphia, PA: Holt-Saunders International. pp. 316–327.
829: This article incorporates text from a publication now in the
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which are external flaps, each with many thin leaf-like membranes.
114:: branchiae) is the zoologists' name for gills (from Ancient Greek
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have modified gills that allow them to live in humid conditions.
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and various mites. A somewhat similar mechanism is used by the
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645:. These thin protuberances on the surface of the body contain
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Sometimes, adults retain these, but they usually disappear at
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Cyclopædia, or an
Universal Dictionary of Arts and Sciences
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ionocytes (formerly known as mitochondrion-rich cells and
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held that fish respired by their gills, but observed that
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Andrews, Chris; Adrian Exell; Neville
Carrington (2003).
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demonstrates that it had internal gills like true fish.
1380:"Plastron respiration in the mite, Platyseius italicus"
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M. b. v. Roberts; Michael Reiss; Grace Monger (2000).
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diffuses into the surrounding water due to its high
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1467:. Updated: 11 June 2010. Retrieved 16 January 2012.
896:Dorit, R. L.; Walker, W. F.; Barnes, R. D. (1991).
1026:(1). Springer Science and Business Media LLC: 24.
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970:"Palatine tonsils—are they branchiogenic organs?"
458:while simultaneously expending energy to excrete
1150:Annales des Sciences Naturelles, Zoologie, Paris
1076:Comparative Physiology of Vertebrate Respiration
626:and moved to the mouth by the beating of cilia.
1341:(Arachnida, Ricinulei) from Central Amazonia".
176:observed that fish had multitudes of openings (
844:(1st ed.). James and John Knapton, et al.
365:gill arch from which projects a cartilaginous
235:that have a highly folded surface to increase
8:
1344:Studies on Neotropical Fauna and Environment
1258:. Discovery Publishing House. p. 180.
223:Gills usually consist of thin filaments of
16:Respiratory organ used by aquatic organisms
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1049:
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291:Freshwater fish gills magnified 400 times
1014:Graham, Anthony; Richardson, Jo (2012).
904:. Saunders College Publishing. pp.
879:. Oxford University Press. 2nd Ed. 1989.
942:. London, UK: Nelson. pp. 164–165.
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889:
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231:(plates), branches, or slender, tufted
1417:Pfingstl, Tobias (30 September 2017).
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299:typically develop in the walls of the
242:A high surface area is crucial to the
1333:Joachim Adis, Benjamin Messner &
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1158:
1079:. Harvard University Press. pp.
750:, as well as at least one species of
254:does, and it diffuses more slowly. A
98:. The gills of some species, such as
34:are visibly exposed as a result of a
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682:, the wall of the caudal end of the
121:With the exception of some aquatic
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86:organisms use to extract dissolved
614:Many marine invertebrates such as
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188:was of another opinion. The word
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426:The red gills inside a detached
384:, lies in the back of the first
51:
1462:Fish Dissection - Gills exposed
1312:. Nelson Thornes. p. 139.
1231:. APH Publishing. p. 269.
1309:Biology: A Functional Approach
1073:Hughes, George Morgan (1963).
968:Slípka, J. (1 December 2003).
1:
1124:; Parsons, Thomas S. (1977).
986:10.1016/S0531-5131(03)01403-1
974:International Congress Series
106:to the external environment.
1436:10.24349/acarologia/20174197
1404:10.1016/0022-1910(71)90184-3
1384:Journal of Insect Physiology
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1282:Biology of Horseshoe Crabs
1183:10.1152/physrev.00050.2003
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258:of air contains about 275
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877:Oxford English Dictionary
430:head (viewed from behind)
359:Broadnose sevengill shark
1365:10.1076/snfe.34.1.1.8915
1306:Roberts, M.B.V. (1986).
1285:. サイエンスハウス. p. 91.
1152:, 13e Série 19: pp 1–42.
781:Artificial gills (human)
570:Pleurobranchaea meckelii
489:, as does the primitive
207:(in singular, meaning a
380:A smaller opening, the
305:countercurrent exchange
276:countercurrent exchange
157:marine animals such as
1279:Sekiguchi, K. (1988).
1255:Text Book of Crustacea
1252:Saxena, Amita (2005).
1122:Romer, Alfred Sherwood
1033:10.1186/2041-9139-3-24
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661:Caribbean hermit crabs
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396:to the ear opening in
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30:The red gills of this
1378:Hinton, H.E. (1971).
1171:Physiological Reviews
955:Manual Of Fish Health
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651:water vascular system
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388:. This bears a small
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201:, "gills", plural of
149:, insects, fish, and
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21:Gill (disambiguation)
715:in atmospheric air.
125:, the filaments and
19:For other uses, see
1396:1971JInsP..17.1185H
1357:1999SNFE...34....1A
1228:Teaching of Biology
1126:The Vertebrate Body
776:Aquatic respiration
764:hydrophilid beetles
752:ricinuleid arachnid
629:Respiration in the
1503:Respiratory system
1339:Cryptocellus adisi
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398:higher vertebrates
346:Cartilaginous fish
321:parathyroid glands
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1488:Arthropod anatomy
1465:Australian Museum
1319:978-0-17-448019-8
1292:978-4-915572-25-8
1265:978-81-8356-016-0
1238:978-81-7648-524-1
1090:978-0-674-15250-2
915:978-0-03-030504-7
857:Missing or empty
850:cite encyclopedia
836:Chambers, Ephraim
485:larvae also have
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100:hermit crabs
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999:18 February
725:respiration
709:hydrophobic
647:diverticula
639:sea urchins
631:echinoderms
582:Crustaceans
512:alpine newt
464:Na/K-ATPase
404:the mouth.
375:gill rakers
297:vertebrates
283:Vertebrates
268:fresh water
256:cubic meter
163:mudskippers
147:crustaceans
32:common carp
1477:Categories
1423:Acarologia
1351:(1): 1–9.
812:References
796:Gill raker
738:, aquatic
721:solubility
609:book gills
527:amphibians
502:Amphibians
495:Polypterus
451:osmolarity
394:homologous
151:amphibians
82:that many
1191:0031-9333
1042:2041-9139
994:0531-5131
980:: 71–74.
801:Gill slit
791:Fish gill
786:Book lung
744:true bugs
705:spiracles
694:Plastrons
680:dragonfly
633:(such as
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575:ctenidium
456:sea water
436:bony fish
418:Bony fish
412:operculum
408:Chimaeras
386:gill slit
337:Fish gill
233:processes
186:Aristotle
36:gill flap
1445:90340235
1199:15618479
1060:23020903
770:See also
700:plastron
673:tracheal
635:starfish
586:molluscs
565:sea slug
540:mudpuppy
538:and the
523:Tadpoles
483:Lungfish
474:Lampreys
447:teleosts
382:spiracle
367:gill ray
229:lamellae
215:Function
204:βράγχιον
190:branchia
178:foramina
143:mollusks
127:lamellae
108:Branchia
1392:Bibcode
1353:Bibcode
1051:3564725
1020:EvoDevo
906:273–276
900:Zoology
833::
740:weevils
736:Elmidae
732:beetles
649:of the
643:papulae
478:hagfish
445:Marine
301:pharynx
198:βράγχια
169:History
123:insects
116:βράγχια
84:aquatic
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677:larval
371:septum
351:Sharks
225:tissue
88:oxygen
1441:S2CID
712:setae
624:mucus
607:have
592:bear
266:. In
260:grams
250:than
194:Greek
174:Galen
159:crabs
131:blood
92:water
90:from
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1314:ISBN
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1195:PMID
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1085:ISBN
1083:–9.
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1038:ISSN
1001:2022
990:ISSN
978:1257
910:ISBN
863:help
806:Lung
671:are
637:and
618:are
476:and
460:salt
428:tuna
355:rays
353:and
331:Fish
161:and
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1431:doi
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