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Gill

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

A gill (/ɡɪl/ (listen)) is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete 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. Branchia (pl. branchiae) is the zoologists' name for gills (from Ancient Greek βράγχια).

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. Semiterrestrial marine animals such as crabs and mudskippers have gill chambers in which they store water, enabling them to use the dissolved oxygen when they are on land.

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Aquatic ecosystem

Aquatic ecosystem

An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms—aquatic life—that are dependent on each other and on their environment. The two main types of aquatic ecosystems are marine ecosystems and freshwater ecosystems. Freshwater ecosystems may be lentic ; lotic ; and wetlands.

Oxygen

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 well as with other compounds. Oxygen is Earth's most abundant element, and after hydrogen and helium, it is the third-most abundant element in the universe. At standard temperature and pressure, two atoms of the element bind to form dioxygen, a colorless and odorless diatomic gas with the formula O2. Diatomic oxygen gas currently constitutes 20.95% of the Earth's atmosphere, though this has changed considerably over long periods of time. Oxygen makes up almost half of the Earth's crust in the form of oxides.

Carbon dioxide

Carbon dioxide

Carbon dioxide (chemical formula CO2) 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, and as the source of available carbon in the carbon cycle, atmospheric CO2 is the primary carbon source for life on Earth. In the air, carbon dioxide is transparent to visible light but absorbs infrared radiation, acting as a greenhouse gas. Carbon dioxide is soluble in water and is found in groundwater, lakes, ice caps, and seawater. When carbon dioxide dissolves in water, it forms carbonate and mainly bicarbonate (HCO−3), which causes ocean acidification as atmospheric CO2 levels increase.

Hermit crab

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 asymmetric abdomen concealed by a snug-fitting shell. Hermit crabs' soft (non-calcified) abdominal exoskeleton means they must occupy shelter produced by other organisms or risk being defenseless.

Surface area

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 length of one-dimensional curves, or of the surface area for polyhedra, for which the surface area is the sum of the areas of its faces. Smooth surfaces, such as a sphere, are assigned surface area using their representation as parametric surfaces. This definition of surface area is based on methods of infinitesimal calculus and involves partial derivatives and double integration.

Insect

Insect

Insects 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, three pairs of jointed legs, compound eyes and one pair of antennae. Their blood is not totally contained in vessels; some circulates in an open cavity known as the haemocoel. Insects are the most diverse group of animals; they include more than a million described species and represent more than half of all known living organisms. The total number of extant species is estimated at between six and ten million; potentially over 90% of the animal life forms on Earth are insects. Insects may be found in nearly all environments, although only a small number of species reside in the oceans, which are dominated by another arthropod group, crustaceans, which recent research has indicated insects are nested within.

Blood

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 circulatory system is also known as peripheral blood, and the blood cells it carries, peripheral blood cells.

Crustacean

Crustacean

Crustaceans 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 can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Amphibian

Amphibian

Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arboreal or freshwater aquatic ecosystems. Thus amphibians typically start out as larvae living in water, but some species have developed behavioural adaptations to bypass this.

Semiaquatic

Semiaquatic

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.

Crab

Crab

Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen), usually hidden entirely under the thorax. They live in all the world's oceans, in freshwater, and on land, are generally covered with a thick exoskeleton, and have a single pair of pincers. They first appeared during the Jurassic Period.

Mudskipper

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 jumping, and the ability to survive prolonged periods of time both in and out of water.

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.[1] The word branchia comes from the Greek βράγχια, "gills", plural of βράγχιον (in singular, meaning a fin).[2]

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Galen

Galen

Aelius Galenus or Claudius Galenus, often Anglicized as Galen or Galen of Pergamon, was a Greek physician, surgeon and philosopher with Roman citizenship. Considered to be one of the most accomplished of all medical researchers of antiquity, Galen influenced the development of various scientific disciplines, including anatomy, physiology, pathology, pharmacology, and neurology, as well as philosophy and logic.

Pliny the Elder

Pliny the Elder

Gaius Plinius Secundus, 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 Naturalis Historia, which became an editorial model for encyclopedias. He spent most of his spare time studying, writing, and investigating natural and geographic phenomena in the field.

Aristotle

Aristotle

Aristotle was an Ancient Greek philosopher and polymath. His writings cover a broad range of subjects including physics, biology, zoology, metaphysics, logic, ethics, aesthetics, poetry, drama, music, rhetoric, psychology, linguistics, economics, politics, meteorology, geology, and government. As the founder of the Peripatetic school of philosophy in the Lyceum in Athens, he began the wider Aristotelian tradition that followed, which set the groundwork for the development of modern science.

Greek language

Greek language

Greek is an independent branch of the Indo-European family of languages, native to Greece, Cyprus, southern Italy, southern Albania, and other regions of the Balkans, the Black Sea coast, Asia Minor, and the Eastern Mediterranean. It has the longest documented history of any Indo-European language, spanning at least 3,400 years of written records. Its writing system is the Greek alphabet, which has been used for approximately 2,800 years; previously, Greek was recorded in writing systems such as Linear B and the Cypriot syllabary. The alphabet arose from the Phoenician script and was in turn the basis of the Latin, Cyrillic, Armenian, Coptic, Gothic, and many other writing systems.

Fin

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. Fins are also used to increase surface areas for heat transfer purposes, or simply as ornamentation.

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.[3]

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.[3]

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. The concentration of oxygen in water is lower than in air and it diffuses more slowly. In fresh water, the dissolved oxygen content is approximately 8 cm3/L compared to that of air which is 210 cm3/L.[4] Water is 777 times more dense than air and is 100 times more viscous.[4] Oxygen has a diffusion rate in air 10,000 times greater than in water.[4] The use of sac-like lungs to remove oxygen from water would not be efficient enough to sustain life.[4] 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, which is what happens when a fish is taken out of water."[4]

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.[3]

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Tissue (biology)

Tissue (biology)

In biology, tissue is a historically derived biological organizational level between cells and a complete organ. A tissue is therefore often thought of as assembly of similar cells and their extracellular matrix from the same origin that together carry out a specific function. Organs are then formed by the functional grouping together of multiple tissues.

Lamella (surface anatomy)

Lamella (surface anatomy)

In surface anatomy, a lamella is a thin plate-like structure, often one amongst many lamellae very close to one another, with open space between. Aside from respiratory organs, they appear in other biological roles including filter feeding and the traction surfaces of geckos.

Process (anatomy)

Process (anatomy)

In anatomy, a process is a projection or outgrowth of tissue from a larger body. For instance, in a vertebra, a process may serve for muscle attachment and leverage, or to fit, with another vertebra. The word is used even at the microanatomic level, where cells can have processes such as cilia or pedicels. Depending on the tissue, processes may also be called by other terms, such as apophysis, tubercle, or protuberance.

Surface area

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 length of one-dimensional curves, or of the surface area for polyhedra, for which the surface area is the sum of the areas of its faces. Smooth surfaces, such as a sphere, are assigned surface area using their representation as parametric surfaces. This definition of surface area is based on methods of infinitesimal calculus and involves partial derivatives and double integration.

Gas exchange

Gas exchange

Gas exchange is the physical process by which gases move passively by 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 liquid, a gas-permeable membrane, or a biological membrane that forms the boundary between an organism and its extracellular environment.

Gram

Gram

The gram is a unit of mass in the International System of Units (SI) equal to one one thousandth of a kilogram.

Fresh water

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 non-salty mineral-rich waters such as chalybeate springs. Fresh water may encompass frozen and meltwater in ice sheets, ice caps, glaciers, snowfields and icebergs, natural precipitations such as rainfall, snowfall, hail/sleet and graupel, and surface runoffs that form inland bodies of water such as wetlands, ponds, lakes, rivers, streams, as well as groundwater contained in aquifers, subterranean rivers and lakes. Fresh water is the water resource that is of the most and immediate use to humans.

Countercurrent exchange

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 other. The flowing bodies can be liquids, gases, or even solid powders, or any combination of those. For example, in a distillation column, the vapors bubble up through the downward flowing liquid while exchanging both heat and mass.

Vertebrates

Freshwater fish gills magnified 400 times
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.[5]

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,[6] parathyroid glands, as well as many other structures derived from the embryonic branchial pouches.[7]

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.[8]

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 pairs and the Broadnose sevengill shark being the only cartilaginous fish exceeding this number. Adjacent slits are separated by a cartilaginous gill arch from which projects a cartilaginous gill ray. This gill ray is the support for the sheet-like interbranchial septum, which the individual lamellae of the gills lie on either side of. The base of the arch may also support gill rakers, projections into the pharyngeal cavity that help to prevent large pieces of debris from damaging the delicate gills.[9]

A smaller opening, the spiracle, lies in the back 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.[9] The spiracle is thought to be homologous to the ear opening in higher vertebrates.[10]

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.[9]

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.[9]

Bony fish

The red gills inside a detached tuna head (viewed from behind)
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 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.[9]

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.[9]

Marine teleosts 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 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 while simultaneously expending energy to excrete salt through the Na+/K+-ATPase ionocytes (formerly known as mitochondrion-rich cells and chloride cells).[11] 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.[9][11]

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 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.[9] Lungfish larvae also have external gills, as does the primitive ray-finned fish Polypterus, though the latter has a structure different from amphibians.[9]

Amphibians

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

Tadpoles of amphibians 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. Examples of salamanders that retain their external gills upon reaching adulthood are the olm and the mudpuppy.

Still, some extinct tetrapod groups did retain true gills. A study on Archegosaurus demonstrates that it had internal gills like true fish.[12]

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Pharynx

Pharynx

The pharynx is the part of the throat behind the mouth and nasal cavity, and above the esophagus and trachea. It is found in vertebrates and invertebrates, though its structure varies across species. The pharynx carries food and air to the esophagus and larynx respectively. The flap of cartilage called the epiglottis stops food from entering the larynx.

Countercurrent exchange

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 other. The flowing bodies can be liquids, gases, or even solid powders, or any combination of those. For example, in a distillation column, the vapors bubble up through the downward flowing liquid while exchanging both heat and mass.

Fish gill

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 like short threads, protein structures called filaments. These filaments have many functions including the transfer of ions and water, as well as the exchange of oxygen, carbon dioxide, acids and ammonia. Each filament contains a capillary network that provides a large surface area for exchanging oxygen and carbon dioxide.

Shark

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 Selachimorpha and are the sister group to the Batoidea. Some sources extend the term "shark" as an informal category including extinct members of Chondrichthyes with a shark-like morphology, such as hybodonts and xenacanths. Shark-like chondrichthyans such as Cladoselache and Doliodus first appeared in the Devonian Period, though some fossilized chondrichthyan-like scales are as old as the Late Ordovician. The oldest modern sharks (selachians) are known from the Early Jurassic, about 200 Ma.

Broadnose sevengill shark

Broadnose sevengill shark

The broadnose sevengill shark 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 the exception of the members of the order Hexanchiformes and the sixgill sawshark. This shark has a large, thick body, with a broad head and blunt snout. The top jaw has jagged, cusped teeth and the bottom jaw has comb-shaped teeth. Its single dorsal fin is set far back along the spine towards the caudal fin, and is behind the pelvic fins. In this shark the upper caudal fin is much longer than the lower, and is slightly notched near the tip. Like many sharks, this sevengill is counter-shaded. Its dorsal surface is silver-gray to brown in order to blend with the dark water and substrate when viewed from above. In counter to this, its ventral surface is very pale, blending with the sunlit water when viewed from below. The body and fins are covered in a scattering of small black & white spots. In juveniles, their fins often have white margins.

Cartilage

Cartilage

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 the bronchial tubes, and the intervertebral discs. In other taxa, such as chondrichthyans, but also in cyclostomes, it may constitute a much greater proportion of the skeleton. It is not as hard and rigid as bone, but it is much stiffer and much less flexible than muscle. The matrix of cartilage is made up of glycosaminoglycans, proteoglycans, collagen fibers and, sometimes, elastin.

Septum

Septum

In biology, a septum 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.

Gill raker

Gill raker

Gill rakers in fish are bony or cartilaginous processes that project from the branchial 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 used for gas exchange. Rakers are usually present in two rows, projecting from both the anterior and posterior side of each gill arch. Rakers are widely varied in number, spacing, and form. By preventing food particles from exiting the spaces between the gill arches, they enable the retention of food particles in filter feeders.

Gill slit

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 contrast, bony fishes have a single outer bony gill covering called an operculum.

Homology (biology)

Homology (biology)

In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.

Chimaera

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.

Operculum (fish)

Operculum (fish)

The operculum is a series of bones found in bony fish and chimaeras that serves as a facial support structure and a protective covering for the gills; it is also used for respiration and feeding.

Invertebrates

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

Crustaceans, molluscs, 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 bear parapodia many of which carry gills.[3] Sponges lack specialised respiratory structures, and the whole of the animal acts as a gill as water is drawn through its spongy structure.[13]

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.[14] Horseshoe crabs have book gills which are external flaps, each with many thin leaf-like membranes.[15]

Many marine invertebrates such as bivalve molluscs are filter feeders. 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 and moved to the mouth by the beating of cilia.[16]

Respiration in the echinoderms (such as 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.

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

The gills of aquatic insects are tracheal, 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 larval dragonfly, the wall of the caudal end of the alimentary tract (rectum) 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 spiracles that connect to the tracheal system. The plastron typically consists of dense patches of hydrophobic setae 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 diffuses into the surrounding water due to its high solubility, while oxygen diffuses into the film as the concentration within the film has been reduced by respiration, 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.

The inorganic 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, as well as at least one species of ricinuleid arachnid.[17] A somewhat similar mechanism is used by the diving bell spider, which maintains an underwater bubble that exchanges gas like a plastron. Other diving insects (such as backswimmers, and hydrophilid beetles) may carry trapped air bubbles, but deplete the oxygen more quickly, and thus need constant replenishment.

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Ctenidium (mollusc)

Ctenidium (mollusc)

A ctenidium is a respiratory organ or gill which is found in many molluscs. This structure exists in bivalves, cephalopods, Polyplacophorans (chitons), and in aquatic gastropods such as freshwater snails and marine snails. Some aquatic gastropods possess one ctenidium known as monopectinate and others have a pair of ctenidia known as bipectinate.

Crustacean

Crustacean

Crustaceans 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 can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Polychaete

Polychaete

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 made of chitin. More than 10,000 species are described in this class. Common representatives include the lugworm and the sandworm or clam worm Alitta.

Parapodium

Parapodium

In invertebrates, the term parapodium refers to lateral outgrowths or protrusions from the body. Parapodia are predominantly found in annelids, where they are paired, unjointed lateral outgrowths that bear the chaetae. In several groups of sea snails and sea slugs, 'parapodium' refers to lateral fleshy protrusions.

Horseshoe crab

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 arachnids such as spiders, ticks, and scorpions.

Bivalvia

Bivalvia

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, bivalves have no head and they lack some usual molluscan organs, like the radula and the odontophore. The class includes the clams, oysters, cockles, mussels, scallops, and numerous other families that live in saltwater, as well as a number of families that live in freshwater. The majority are filter feeders. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment, where they are relatively safe from predation. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and file shells, can swim. The shipworms bore into wood, clay, or stone and live inside these substances.

Filter feeder

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 feeding are clams, krill, sponges, baleen whales, and many fish. Some birds, such as flamingos and certain species of duck, are also filter feeders. Filter feeders can play an important role in clarifying water, and are therefore considered ecosystem engineers. They are also important in bioaccumulation and, as a result, as indicator organisms.

Mucus

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 a viscous colloid containing inorganic salts, antimicrobial enzymes, immunoglobulins, and glycoproteins such as lactoferrin and mucins, which are produced by goblet cells in the mucous membranes and submucosal glands. Mucus serves to protect epithelial cells in the linings of the respiratory, digestive, and urogenital systems, and structures in the visual and auditory systems from pathogenic fungi, bacteria and viruses. Most of the mucus in the body is produced in the gastrointestinal tract.

Echinoderm

Echinoderm

An echinoderm is any member of the phylum Echinodermata. The adults are recognisable by their radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea lilies or "stone lilies". Adult echinoderms are found on the sea bed at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 7,000 living species, making it the second-largest grouping of deuterostomes, after the chordates. Echinoderms are the largest entirely marine phylum. The first definitive echinoderms appeared near the start of the Cambrian.

Diverticulum

Diverticulum

In medicine or biology, a diverticulum is an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, diverticula are described as being either true or false.

Caribbean hermit crab

Caribbean hermit crab

The Caribbean hermit crab, also known as the soldier crab, West Atlantic crab, tree crab, or purple pincher, is a species of land hermit crab native to the west Atlantic, Belize, southern Florida, Venezuela, and the West Indies.

Aquatic insect

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 cannot compete.

Source: "Gill", Wikipedia, Wikimedia Foundation, (2022, December 7th), https://en.wikipedia.org/wiki/Gill.

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References
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