Gill arches bearing gills in a pike. Gills allow fish to breathe underwater.
Respiratory mechanism in bony fish
The fish draws oxygen-rich water in through the mouth (left). It then pumps it over gills so oxygen enters the bloodstream, and allows oxygen-depleted water to exit through the gill slits (right)
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.[1][2] Each filament contains a capillary network that provides a large surface area for exchanging oxygen and carbon dioxide.
Fish exchange gases by pulling oxygen-rich water through their mouths and pumping it over their gills. Within the gill filaments, capillary blood flows in the opposite direction to the water, causing counter-current exchange. The gills push the oxygen-poor water out through openings in the sides of the pharynx. Some fish, like sharks and lampreys, possess multiple gill openings. However, bony fish have a single gill opening on each side. This opening is hidden beneath a protective bony cover called the operculum.
Juvenile bichirs have external gills, a very primitive feature that they share with larval amphibians.
Previously, the evolution of gills was thought to have occurred through two diverging lines: gills formed from the endoderm, as seen in jawless fish species, or those form by the ectoderm, as seen in jawed fish. However, recent studies on gill formation of the little skate (Leucoraja erinacea) has shown potential evidence supporting the claim that gills from all current fish species have in fact evolved from a common ancestor.[3]
Breathing with gills
Tuna gills inside the head. The head is oriented snout-down with the view looking towards the mouth.
The red gills detached from the tuna head on the left
All basal vertebrates (types of fish) breathe with gills. The gills are carried right behind the head, bordering the posterior margins of a series of openings from the esophagus to the exterior. Each gill is supported by a cartilaginous or bony gill arch.[4] 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 counter-current 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 that it passes over the gills to the outside. The bony fish have three pairs of arches, cartilaginous fish have five to seven pairs, while the primitive jawless fish have seven. The vertebrate ancestor no doubt had more arches, as some of their chordate relatives have more than 50 pairs of gills.[6]
Gills usually consist of thin filaments of tissue, branches, or slender tufted processes that have a highly folded surface to increase surface area. The 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 air and it diffuses more slowly. In a litre of freshwater the oxygen content is 8cm3 per litre compared to 210 in the same volume of air.[7] Water is 777 times more dense than air and is 100 times more viscous.[7] Oxygen has a diffusion rate in air 10,000 times greater than in water.[7] The use of sac-like lungs to remove oxygen from water would not be efficient enough to sustain life.[7] Rather than using lungs "Gaseous 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."[7]
In bony fish, the gills lie in a branchial chamber covered by a bony operculum (branchia is an Ancient Greek word for gills). 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 that 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.[6]
The gill arches of bony fish typically have no septum, so that 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 a spiracle, the pseudobranch associated with it 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.[6]
Most bony fish have five gills
Fish transfer oxygen from the sea water to their blood using a highly efficient mechanism called countercurrent exchange. Countercurrent exchange means the flow of water over the gills is in the opposite direction to the flow of blood through the capillaries in the lamellae. The effect of this is that the blood flowing in the capillaries always encounters water with a higher oxygen concentration, allowing diffusion to occur all the way along the lamellae. As a result the gills can extract over 80% of the oxygen available in the water.
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).[10] Conversely, freshwater has 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.[6][10]
In some primitive bony fishes and amphibians, the larvae bear external gills, branching off from the gill arches.[11] These are reduced in adulthood, their function taken over by the gills proper in fishes and by lungs in most amphibians. Some amphibians retain the external larval gills in adulthood, the complex internal gill system as seen in fish apparently being irrevocably lost very early in the evolution of tetrapods.[12]
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 or seven pairs. Adjacent slits are separated by a cartilaginous gill arch from which projects a long sheet-like septum, partly supported by a further piece of cartilage called the gill ray. The individual lamellae of the gills lie on either side of the septum. The base of the arch may also support gill rakers, small projecting elements that help to filter food from the water.[6]
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.[6] The spiracle is thought to be homologous to the ear opening in higher vertebrates.[13]
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.[6]
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.[6]
The shared trait of breathing via gills in bony fish and cartilaginous fish is a famous example of symplesiomorphy. Bony fish are more closely related to terrestrial vertebrates, which evolved out of a clade of bony fishes that breathe through their skin or lungs, than they are to the sharks, rays, and the other cartilaginous fish. Their kind of gill respiration is shared by the "fishes" because it was present in their common ancestor and lost in the other living vertebrates. But based on this shared trait, we cannot infer that bony fish are more closely related to sharks and rays than they are to terrestrial vertebrates.[14]
Lampreys and hagfish
Outline of a hagfish, showing above the two ventral openings (h) by which the water escapes from the gills, and in the dissection below the spherical pouches which contain the gills
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. In adult lampreys, a separate respiratory tube develops beneath the pharynx proper, separating food and water from respiration by closing a valve at its anterior end.[6]
Breathing without gills
Some fish can at least partially respire without gills. In some species cutaneous respiration accounts for 5 to 40 per cent of the total respiration, depending on temperature. Cutaneous respiration is more important in species that breathe air, such as mudskippers and reedfish, and in such species can account for nearly half the total respiration.[15]
Fish from multiple groups can live out of the water for extended time periods.
Air breathing fish can be divided into obligate air breathers and facultative air breathers. Obligate air breathers, such as the African lungfish, are obligated to breathe air periodically or they suffocate. Facultative air breathers, such as the catfish Hypostomus plecostomus, only breathe air if they need to and can otherwise rely on their gills for oxygen. Most air breathing fish are facultative air breathers that avoid the energetic cost of rising to the surface and the fitness cost of exposure to surface predators.[16]
Osteichthyes, also known as osteichthyans or commonly referred to as the bony fish, is a diverse superclass of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes and the extinct placoderms and acanthodians, which have endoskeletons primarily composed of cartilage. The vast majority of extant fish are members of Osteichthyes, being an extremely diverse and abundant group consisting of 45 orders, over 435 families and 28,000 species. It is the largest class of vertebrates in existence today, encompassing most aquatic vertebrates, as well as all semi-aquatic and terrestrial vertebrates.
A gill 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 is the zoologists' name for gills.
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.
Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, including the presence of lobed fins with a well-developed internal skeleton. Lungfish represent the closest living relatives of the tetrapods. The mouths of lungfish typically bear tooth plates, which are used to crush hard shelled organisms.
Bichirs and the reedfish comprise Polypteridae, a family of archaic ray-finned fishes and the only family in the order Polypteriformes.
Fish anatomy is the study of the form or morphology of fish. It can be contrasted with fish physiology, which is the study of how the component parts of fish function together in the living fish. In practice, fish anatomy and fish 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 latter dealing with how those components function together in living fish.
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.
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.
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.
Rajiformes is one of the four orders in the clade Batomorphi, often referred to as the superorder Batoidea, flattened cartilaginous fishes related to sharks. Rajiforms are distinguished by the presence of greatly enlarged pectoral fins, which reach as far forward as the sides of the head, with a generally flattened body. The undulatory pectoral fin motion diagnostic to this taxon is known as rajiform locomotion. The eyes and spiracles are located on the upper surface of the head and the gill slits are on the underside of the body. Most species give birth to live young, although some lay eggs enclosed in a horny capsule.
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.
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.
Amphibious fish are fish that are able to leave water for extended periods of time. About 11 distantly related genera of fish are considered amphibious. This suggests that many fish genera independently evolved amphibious traits, a process known as convergent evolution. These fish use a range of methods for land movement, such as lateral undulation, tripod-like walking, and jumping. Many of these methods of locomotion incorporate multiple combinations of pectoral-, pelvic-, and tail-fin movement.
This glossary of ichthyology is a list of definitions of terms and concepts used in ichthyology, the study of fishes.
Shark anatomy differs from that of bony fish in a variety of ways. Variation observed within shark anatomy is a potential result of speciation and habitat variation.
Branchial arches, or gill arches, are a series of paired bony "loops" that support the gills in fish. As gills are the primitive feature of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arches varies between taxa. In jawed fish, the first arch pair develops into the jaw. The second gill arches develop into the hyomandibular complex, which supports the back of the jaw and the front of the gill series. The remaining posterior arches support the gills. In amphibians and reptiles, many pharyngeal arch elements are lost, including the gill arches, resulting in only the oral jaws and a hyoid apparatus remaining. In mammals and birds, the hyoid is simplified further.
Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.
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 latter dealing with how those components function together in the living fish.
Spiracles are openings on the surface of some animals, which usually lead to respiratory systems.
1 2 Evans, David H.; Piermarini, Peter M.; Choe, Keith P. (January 2005). "The Multifunctional Fish Gill: Dominant Site of Gas Exchange, Osmoregulation, Acid-Base Regulation, and Excretion of Nitrogenous Waste". Physiological Reviews. 85 (1): 97–177. doi:10.1152/physrev.00050.2003. ISSN0031-9333. PMID15618479.
↑ Szarski, Henryk (1957). "The Origin of the Larva and Metamorphosis in Amphibia". The American Naturalist. 91 (860). Essex Institute: 287. doi:10.1086/281990. JSTOR2458911. S2CID85231736.
↑ Clack, J. A. (2002): Gaining ground: the origin and evolution of tetrapods. Indiana University Press, Bloomington, Indiana. 369 pp
↑ Laurin M. (1998): The importance of global parsimony and historical bias in understanding tetrapod evolution. Part I-systematics, middle ear evolution, and jaw suspension. Annales des Sciences Naturelles, Zoologie, Paris, 13e Série 19: pp 1-42.
↑ Cracraft, Joel; Donoghue, Michael J. (2004), Assembling the Tree of Life, USA: Oxford University Press, p.367, ISBN0-19-517234-5
↑ Pozdnyakov, S. E. & Gibson, D. I. (2008). Family Didymozoidae Monticelli, 1888. In R. A. Bray, D. I. Gibson & A. Jones (Eds.), Keys to the Trematoda, Vol. 3 (pp. 631-734). London: CAB International and The Natural History Museum.
↑ Justine, JL. (September 2004). "Three new species of Huffmanela Moravec, 1987 (Nematoda: Trichosomoididae) from the gills of marine fish off New Caledonia". Systematic Parasitology. 59 (1): 29–37. doi:10.1023/B:SYPA.0000038442.25230.8b. PMID15318018. S2CID29105973.
Evans, D H; Piermarini, P M; Choe, K P (2005). "The multifunctional fish gill: dominant site of gas exchange, osmoregulation, acid-base regulation, and excretion of nitrogenous waste". Physiological Reviews. 85 (1): 97–177. doi:10.1152/physrev.00050.2003. PMID15618479.
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