Gill

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

A gill ( /ɡɪl/ ( Loudspeaker.svg 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).

Contents

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.

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]

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]

Vertebrates

Freshwater fish gills magnified 400 times FreshwaterFishGill400x7.jpg
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) Tuna Gills in Situ cut.jpg
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 Smooth Newt larva (aka).jpg
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]

Invertebrates

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

See also

Related Research Articles

<span class="mw-page-title-main">Respiratory system</span> Biological system in animals and plants for gas exchange

The respiratory system is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies greatly, depending on the size of the organism, the environment in which it lives and its evolutionary history. In land animals the respiratory surface is internalized as linings of the lungs. Gas exchange in the lungs occurs in millions of small air sacs; in mammals and reptiles these are called alveoli, and in birds they are known as atria. These microscopic air sacs have a very rich blood supply, thus bringing the air into close contact with the blood. These air sacs communicate with the external environment via a system of airways, or hollow tubes, of which the largest is the trachea, which branches in the middle of the chest into the two main bronchi. These enter the lungs where they branch into progressively narrower secondary and tertiary bronchi that branch into numerous smaller tubes, the bronchioles. In birds the bronchioles are termed parabronchi. It is the bronchioles, or parabronchi that generally open into the microscopic alveoli in mammals and atria in birds. Air has to be pumped from the environment into the alveoli or atria by the process of breathing which involves the muscles of respiration.

<span class="mw-page-title-main">Book lung</span> Type of lung commonly found in arachnids

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 connects with the surroundings through a small opening for the purpose of respiration.

<span class="mw-page-title-main">Aquatic respiration</span> Process whereby an aquatic animal obtains oxygen from water

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.

<span class="mw-page-title-main">Aquatic insect</span> Insect that lives in water

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.

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.

<span class="mw-page-title-main">Ostracoderm</span> Armored jawless fish of the Paleozoic

Ostracoderms are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic and thus does not correspond to one evolutionary lineage. However, the term is still used as an informal way of loosely grouping together the armored jawless fishes.

<span class="mw-page-title-main">Lamella (surface anatomy)</span> Anatomical structure

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.

Gill slit Individual opening to a gill

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.

Operculum (fish) Bones in a fish that provide facial support structure and a protective covering for the gills

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.

<span class="mw-page-title-main">Fish</span> Aquatic vertebrates, usually with gills

Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of living fish species are ray-finned fish, belonging to the class Actinopterygii, with around 99% of those being teleosts.

This glossary of ichthyology is a list of definitions of terms and concepts used in ichthyology, the study of fishes.

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Insect physiology includes the physiology and biochemistry of insect organ systems.

Respiratory system of insects

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Fish are exposed to large oxygen fluctuations in their aquatic environment since the inherent properties of water can result in marked spatial and temporal differences in the concentration of oxygen. Fish respond to hypoxia with varied behavioral, physiological, and cellular responses in order to maintain homeostasis and organism function in an oxygen-depleted environment. The biggest challenge fish face when exposed to low oxygen conditions is maintaining metabolic energy balance, as 95% of the oxygen consumed by fish is used for ATP production releasing the chemical energy of nutrients through the mitochondrial electron transport chain. Therefore, hypoxia survival requires a coordinated response to secure more oxygen from the depleted environment and counteract the metabolic consequences of decreased ATP production at the mitochondria. This article is a review of the effects of hypoxia on all aspects of fish, ranging from behavior down to genes.

Fish gill Organ that allows fish to breathe underwater

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.

Fish physiology Scientific study of how the component parts of fish function together in the living fish

Fish physiology is the scientific study of how the component parts of fish function together in the living fish. It can be contrasted with fish anatomy, which is the study of the form or morphology of fishes. In practice, fish anatomy and physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the later dealing with how those components function together in the living fish. For this, at first we need to know about their intestinal morphology.

Cutaneous respiration, or cutaneous gas exchange, is a form of respiration in which gas exchange occurs across the skin or outer integument of an organism rather than gills or lungs. Cutaneous respiration may be the sole method of gas exchange, or may accompany other forms, such as ventilation. Cutaneous respiration occurs in a wide variety of organisms, including insects, amphibians, fish, sea snakes, turtles, and to a lesser extent in mammals.

Spiracle (vertebrates)

Spiracles are openings on the surface of some animals, which usually lead to respiratory systems.

References

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