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A gill () is a respiratory organ that many aquatic organisms use to extract dissolved
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
from
water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a ...
and to excrete
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
. The gills of some species, such as
hermit crab Hermit crabs are anomuran decapod crustaceans of the superfamily Paguroidea that have adapted to occupy empty scavenged mollusc shells to protect their fragile exoskeletons. There are over 800 species of hermit crab, most of which possess an a ...
s, have adapted to allow respiration on land provided they are kept moist. The microscopic structure of a gill presents a large
surface area The surface area of a solid object is a measure of the total area that the surface of the object occupies. The mathematical definition of surface area in the presence of curved surfaces is considerably more involved than the definition of arc ...
to the external environment. Branchia (pl. branchiae) is the zoologists' name for gills (from Ancient Greek ). With the exception of some aquatic
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, the filaments and lamellae (folds) contain
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the c ...
or coelomic fluid, from which gases are exchanged through the thin walls. The blood carries oxygen to other parts of the body.
Carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
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
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s, insects, fish, and
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s.
Semiterrestrial In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
marine animals such as
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s and
mudskipper Mudskippers are any of the 23 extant species of amphibious fish from the subfamily Oxudercinae of the goby family Oxudercidae. They are known for their unusual body shapes, preferences for semiaquatic habitats, limited terrestrial locomotion and ...
s have gill chambers in which they store water, enabling them to use the dissolved oxygen when they are on land.


History

Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be one of ...
observed that fish had multitudes of openings (''foramina''), big enough to admit gases, but too fine to give passage to water.
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic '' ...
held that fish respired by their gills, but observed that
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
was of another opinion. The word ''branchia'' comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, "gills", plural of (in singular, meaning a
fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. 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 a gill or gills. Many invertebrates, and even amphibians, use both the body surface and gills for gaseous exchange. Gills usually consist of thin filaments of tissue,
lamellae Lamella (plural lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * ...
(plates), branches, or slender, tufted processes that have a highly folded surface to increase
surface area The surface area of a solid object is a measure of the total area that the surface of the object occupies. The mathematical definition of surface area in the presence of curved surfaces is considerably more involved than the definition of arc ...
. 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 Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
of aquatic organisms, as water contains only a small fraction of the
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
that
air The atmosphere of Earth is the layer of gases, known collectively as air, retained by Earth's gravity that surrounds the planet and forms its planetary atmosphere. The atmosphere of Earth protects life on Earth by creating pressure allowing for ...
does. A
cubic meter The cubic metre (in Commonwealth English and international spelling as used by the International Bureau of Weights and Measures) or cubic meter (in American English) is the unit of volume in the International System of Units (SI). Its symbol is m ...
of air contains about 250
gram The gram (originally gramme; SI unit symbol g) is a Physical unit, unit of mass in the International System of Units (SI) equal to one one thousandth of a kilogram. Originally defined as of 1795 as "the absolute weight of a volume of pure wate ...
s of oxygen at
STP STP may refer to: Places * São Tomé and Príncipe (ISO 3166-1 alpha-3 code, IOC country code, and FIFA country code STP) * London St Pancras (Domestic) railway station (National Rail code STP) * St. Paul Downtown Airport (IATA airport code S ...
. The concentration of oxygen in water is lower than in air and it diffuses more slowly. In
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
, the dissolved oxygen content is approximately 8 cm3/L compared to that 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, " seous 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." 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 Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each ...
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
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s typically develop in the walls of the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
, along a series of gill slits opening to the exterior. Most species employ a
countercurrent exchange Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each ...
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 gland The thymus is a specialized primary lymphoid organ of the immune system. Within the thymus, thymus cell lymphocytes or ''T cells'' mature. T cells are critical to the adaptive immune system, where the body adapts to specific foreign invaders. ...
s,
parathyroid glands Parathyroid glands are small endocrine glands in the neck of humans and other tetrapods. Humans usually have four parathyroid glands, located on the back of the thyroid gland in variable locations. The parathyroid gland produces and secretes pa ...
, as well as many other structures derived from the embryonic
branchial pouch In the embryonic development of vertebrates, pharyngeal pouches form on the endodermal side between the pharyngeal arches. The pharyngeal grooves (or clefts) form the lateral ectodermal surface of the neck region to separate the arches. The pouch ...
es.


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.


Cartilaginous fish

Shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s and
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six pairs and the
Broadnose sevengill shark The broadnose sevengill shark (''Notorynchus cepedianus'') is the only extant member of the genus ''Notorynchus'', in the family Hexanchidae. It is recognizable because of its seven gill slits, while most shark species have five gill slits, with ...
being the only cartilaginous fish exceeding this number. Adjacent slits are separated by a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...
gill arch from which projects a cartilaginous gill ray. This gill ray is the support for the sheet-like interbranchial
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatri ...
, which the individual lamellae of the gills lie on either side of. The base of the arch may also support
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of the ...
s, projections into the pharyngeal cavity that help to prevent large pieces of debris from damaging the delicate gills. A smaller opening, the
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
, lies in the back of the first
gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
. This bears a small pseudobranch that resembles a gill in structure, but only receives blood already oxygenated by the true gills. The spiracle is thought to be homologous to the ear opening in
higher vertebrates Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
. 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.
Chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
s 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.


Bony fish

In
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
, 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. 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 base of the operculum. This is, however, often greatly reduced, consisting of a small mass of cells without any remaining gill-like structure. Marine
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s 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
osmolarity Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). The osmolarity of a solution is usually expressed as Osm/L ...
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
sea water Seawater, or salt water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has approx ...
while simultaneously expending energy to excrete
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
through the Na+/K+-ATPase ionocytes (formerly known as mitochondrion-rich cells and
chloride cell An ionocyte (formerly called a chloride cell) is a mitochondrion-rich cell within ionoregulatory organs of animals, such as teleost fish gill, insect Malpighian tubules, crustacean gills, antennal glands and maxillary glands, and copepod Crusalis ...
s). 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.
Lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s and
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
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 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.
Lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
larvae also have
external gills External gills are the gills of an animal, most typically an amphibian, that are exposed to the environment, rather than set inside the pharynx and covered by gill slits, as they are in most fishes. Instead, the respiratory organs are set on a frill ...
, as does the primitive
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or hor ...
''
Polypterus ''Polypterus'' is a genus of freshwater fish in the bichir family (Polypteridae) of order Polypteriformes. The type species is the Nile bichir (''P. bichir''). Fish in this genus live in various areas in Africa. ''Polypterus'' is the only know ...
'', though the latter has a structure different from amphibians.


Amphibians

Tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found i ...
s of
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s 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. Sometimes, adults retain these, but they usually disappear at
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
. Examples of salamanders that retain their external gills upon reaching adulthood are the
olm The olm or proteus (''Proteus anguinus'') is an aquatic salamander in the family Proteidae, the only exclusively cave-dwelling chordate species found in Europe. In contrast to most amphibians, it is entirely aquatic, eating, sleeping, and br ...
and the
mudpuppy ''Necturus'' is a genus of aquatic salamanders native to the eastern United States and Canada. They are commonly known as waterdogs and mudpuppies. The common mudpuppy ''(N. maculosus)'' is probably the best-known species – as an amphibian wi ...
. Still, some extinct tetrapod groups did retain true gills. A study on ''
Archegosaurus ''Archegosaurus'' is a genus of temnospondyl amphibian which lived during the Asselian to Wuchiapingian stages of the Permian, around 299-253 million years ago. The remains of this animal, consisting of at least 90 partial skeletons (mostly skull ...
'' demonstrates that it had internal gills like true fish.


Invertebrates

Crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s, 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
polychaete worms Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are m ...
bear
parapodia In invertebrates, the term parapodium ( Gr. ''para'', beyond or beside + ''podia'', feet; plural: parapodia) refers to lateral outgrowths or protrusions from the body. Parapodia are predominantly found in annelids, where they are paired, unjointed ...
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. 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 gill chamber.
Horseshoe crab Horseshoe crabs are marine and brackish water arthropods of the family Limulidae and the only living members of the order Xiphosura. Despite their name, they are not true crabs or crustaceans: they are chelicerates, most closely related to arachn ...
s have book gills which are external flaps, each with many thin leaf-like membranes. Many marine invertebrates such as
bivalve molluscs Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
are
filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s. 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
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is ...
and moved to the mouth by the beating of cilia. Respiration in the
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s (such as
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
and
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) of ...
s) is carried out using a very primitive version of gills called
papulae Papulae (sing. papula; also occasionally papulla, papullae), also known as dermal branchiae or skin gills, are projections of the coelom of Asteroidea that serve in respiration and waste removal. Papulae are soft, covered externally with the epi ...
. These thin protuberances on the surface of the body contain
diverticula In medicine or biology, a diverticulum is an outpouching of a hollow (or a fluid-filled) structure in the body. Depending upon which layers of the structure are involved, diverticula are described as being either true or false. In medicine, t ...
of the
water vascular system The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms mov ...
. The gills of
aquatic insect Aquatic insects or water insects live some portion of their life cycle in the water. They feed in the same ways as other insects. Some ''diving'' insects, such as predatory diving beetles, can hunt for food underwater where land-living insects c ...
s are
tracheal The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
, 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
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l
dragonfly A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
, the wall of the caudal end of the
alimentary tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
(
rectum The rectum is the final straight portion of the large intestine in humans and some other mammals, and the Gastrointestinal tract, gut in others. The adult human rectum is about long, and begins at the rectosigmoid junction (the end of the s ...
) is richly supplied with tracheae as a rectal gill, and water pumped into and out of the rectum provides oxygen to the closed tracheae.


Plastrons

A plastron 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
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
s that connect to the tracheal system. The plastron typically consists of dense patches of
hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, th ...
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
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 in atmospheric air.
Carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
diffuses into the surrounding water due to its high
solubility In chemistry, solubility is the ability of a substance, the solute, to form a solution with another substance, the solvent. Insolubility is the opposite property, the inability of the solute to form such a solution. The extent of the solubil ...
, while oxygen diffuses into the film as the concentration within the film has been reduced by
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellula ...
, 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 Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a s ...
, so many such insects in still water actively direct a flow of water over their bodies. The inorganic gill mechanism allows aquatic insects with plastrons to remain constantly submerged. Examples include many
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s in the family
Elmidae Elmidae, commonly known as riffle beetles, is a family (biology), family of beetles in the superfamily Byrrhoidea described by John Curtis (entomologist), John Curtis in 1830. Both adults and larvae are usually aquatic, living under rocks in fas ...
, aquatic
weevil Weevils are beetles belonging to the Taxonomic rank, superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and Herbivore, herbivorous. Approximately 97,000 species of weevils are known. They b ...
s, and
true bug Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to around ...
s in the family
Aphelocheiridae ''Aphelocheirus'' is a genus of true bug Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shiel ...
, as well as at least one species of ricinuleid arachnid. A somewhat similar mechanism is used by the
diving bell spider The diving bell spider or water spider (''Argyroneta aquatica'') is the only species of spider known to live almost entirely under water. It is the only member of the genus ''Argyroneta''. When out of the water, the spider ranges in colour from m ...
, which maintains an underwater bubble that exchanges gas like a plastron. Other diving insects (such as
backswimmer Notonectidae is a cosmopolitan family of aquatic insects in the order Hemiptera, commonly called backswimmers because they swim "upside down" (inverted). They are all predators and typically range from in length. They are similar in appearance t ...
s, and hydrophilid beetles) may carry trapped air bubbles, but deplete the oxygen more quickly, and thus need constant replenishment.


See also

*
Aquatic respiration Aquatic respiration is the process whereby an aquatic organism exchanges respiratory gases with water, obtaining oxygen from oxygen dissolved in water and excreting carbon dioxide and some other metabolic waste products into the water. Unicel ...
*
Artificial gills (human) Artificial gills are unproven conceptualised devices to allow a human to be able to take in oxygen from surrounding water. This is speculative technology that has not been demonstrated in a documented fashion. Natural gills work because nearly a ...
*
Book lung A book lung is a type of respiration organ used for atmospheric gas exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open ventral abdominal, air-filled cavity (atrium) and conn ...
*
Fish gill Fish gills are organs that allow fish to breathe underwater. Most fish exchange gases like oxygen and carbon dioxide using gills that are protected under gill covers (operculum) on both sides of the pharynx (throat). Gills are tissues that are ...
*
Gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of the ...
*
Gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
*
Lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...


References


External links


Fish Dissection - Gills exposed
''Australian Museum''. Updated: 11 June 2010. Retrieved 16 January 2012. {{Use dmy dates, date=May 2017 Animal anatomy Arthropod anatomy Fish anatomy Organs (anatomy) Respiratory system