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Fish

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For other uses, see Fish (disambiguation). Fishes A giant grouper at the Georgia Aquarium, seen swimming among schools of other fish The ornate lionfish as seen from a head-on view Scientific classification Kingdom: Animalia Phylum: Chordata (unranked) Craniata Included groups Jawless fishes Armoured fishes (extinct) Cartilaginous fishes Ray-finned fishes Lobe-finned fishes Excluded groups

Tetrapods

A fish is any aquatic vertebrate animal that is covered with scales, and equipped with two sets of paired fins and several unpaired fins. Most fish are ectothermic (or cold-blooded). Fish are abundant in most bodies of water. Fish are found in high mountain streams (e.g., char and gudgeon) and in the deepest ocean depths (e.g., gulpers and anglerfish).

Food prepared from fish is also called fish, and is an important human food source. Commercial and subsistence fishers ``hunt`` them in wild fisheries (see fishing) or ``farm`` them in ponds or in cages in the ocean (see aquaculture). They are also caught by recreational fishers and raised by fishkeepers, and are exhibited in public aquaria. Fish have had a role in culture through the ages, serving as deities, religious symbols, and as the subjects of art, books and movies.

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Diversity of fish

Main article: Diversity of fish

The term ``fish`` most precisely describes any non-tetrapod craniate (i.e. an animal with a skull and in most cases a backbone) that has gills throughout life and whose limbs, if any, are in the shape of fins.[1] Unlike groupings such as birds or mammals, fish are not a single clade but a paraphyletic collection of taxa, including hagfishes, lampreys, sharks and rays, ray-finned fishes, coelacanths, and lungfishes.[2][3]

A typical fish is ectothermic, has a streamlined body for rapid swimming, extracts oxygen from water using gills or uses an accessory breathing organ to breathe atmospheric oxygen, has two sets of paired fins, usually one or two (rarely three) dorsal fins, an anal fin, and a tail fin, has jaws, has skin that is usually covered with scales, and lays eggs.

Fish come in many shapes and sizes. This is a sea dragon, a close relative of the seahorse. Their leaf-like appendages enable them to blend in with floating seaweed.

Each criterion has exceptions. Tuna, swordfish, and some species of sharks show some warm-blooded adaptations—they can heat their bodies significantly above ambient water temperature.[2] Streamlining and swimming performance varies from fish such as tuna, salmon, and jacks that can cover 10–20 body-lengths per second to species such as eels and rays that swim no more than 0.5 body-lengths per second.[2] Many groups of freshwater fish extract oxygen from the air as well as from the water using a variety of different structures. Lungfish have paired lungs similar to those of tetrapods, gouramis have a structure called the labyrinth organ that performs a similar function, while many catfish, such as Corydoras extract oxygen via the intestine or stomach.[4] Body shape and the arrangement of the fins is highly variable, covering such seemingly un-fishlike forms as seahorses, pufferfish, anglerfish, and gulpers. Similarly, the surface of the skin may be naked (as in moray eels), or covered with scales of a variety of different types usually defined as placoid (typical of sharks and rays), cosmoid (fossil lungfishes and coelacanths), ganoid (various fossil fishes but also living gars and bichirs), cycloid, and ctenoid (these last two are found on most bony fish).[5] There are even fishes that live mostly on land. Mudskippers feed and interact with one another on mudflats and go underwater to hide in their burrows.[6] The catfish Phreatobius cisternarum lives in underground, phreatic habitats, and a relative lives in waterlogged leaf litter.[7][8]

Fish range in size from the huge 16 metres (52 ft) whale shark to the tiny 8 millimetres (0.31 in) stout infantfish.

Many types of aquatic animals commonly referred to as ``fish`` are not fish in the sense given above; examples include shellfish, cuttlefish, starfish, crayfish and jellyfish. In earlier times, even biologists did not make a distinction – sixteenth century natural historians classified also seals, whales, amphibians, crocodiles, even hippopotamuses, as well as a host of aquatic invertebrates, as fish.[9] In some contexts, especially in aquaculture, the true fish are referred to as finfish (or fin fish) to distinguish them from these other animals.

Taxonomy

Fish are a paraphyletic group: that is, any clade containing all fish also contains the tetrapods, which are not fish. For this reason, groups such as the ``Class Pisces`` seen in older reference works are no longer used in formal classifications.

Fish are classified into the following major groups:

Subclass Pteraspidomorphi (early jawless fish) Class Thelodonti Class Anaspida (unranked) Cephalaspidomorphi (early jawless fish) (unranked) Hyperoartia or Petromyzontida Petromyzontidae (lampreys) Class Galeaspida Class Pituriaspida Class Osteostraci Infraphylum Gnathostomata (jawed vertebrates) Class Placodermi (armoured fishes, extinct) Class Chondrichthyes (cartilaginous fish) Class Acanthodii (spiny sharks, extinct) Superclass Osteichthyes (bony fish) Class Actinopterygii (ray-finned fish) Subclass Chondrostei Order Acipenseriformes (sturgeons and paddlefishes) Order Polypteriformes (reedfishes and bichirs). Subclass Neopterygii Infraclass Holostei (gars and bowfins) Infraclass Teleostei (many orders of common fishes) Class Sarcopterygii (lobe-finned fish) Subclass Coelacanthimorpha (coelacanths) Subclass Dipnoi (lungfish)

Some palaeontologists contend that because Conodonta are chordates, they are primitive fish. For a fuller treatment of this taxonomy, see the vertebrate article.

The various fish groups account for more than half of vertebrate species. There are almost 28,000 known extant species, of which almost 27,000 are bony fish, with 970 sharks, rays, and chimeras and about 108 hagfishes and lampreys.[10] A third of these species fall within the nine largest families; from largest to smallest, these families are Cyprinidae, Gobiidae, Cichlidae, Characidae, Loricariidae, Balitoridae, Serranidae, Labridae, and Scorpaenidae. About 64 families are monotypic, containing only one species. The final total of extant species may grow to exceed 32,500.[11]

Anatomy

Main article: Fish anatomy The anatomy of Lampanyctodes hectoris (1) – operculum (gill cover), (2) – lateral line, (3) – dorsal fin, (4) – fat fin, (5) – caudal peduncle, (6) – caudal fin, (7) – anal fin, (8) – photophores, (9) – pelvic fins (paired), (10) – pectoral fins (paired)

Respiration

Most fish exchange gases using gills on either side of the pharynx. Gills consist of threadlike structures called filaments. 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. In some fishes, 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 fishes, like sharks and lampreys, possess multiple gill openings. However, most fishes have a single gill opening on each side. This opening is hidden beneath a protective bony cover called an operculum.

Juvenile bichirs have external gills, a very primitive feature that they share with larval amphibians.

Swim bladder of a Rudd (Scardinius erythrophthalmus)

Many fish can breathe air via a variety of mechanisms. The skin of anguillid eels may absorb oxygen. The buccal cavity of the electric eel may breathe air. Catfishes of the families Loricariidae, Callichthyidae, and Scoloplacidae absorb air through their digestive tracts.[12] Lungfish and bichirs have paired lungs similar to those of tetrapods and must surface to gulp fresh air through the mouth and pass spent air out through the gills. Gar and bowfin have a vascularized swim bladder that functions in the same way. Loaches, trahiras, and many catfish breathe by passing air through the gut. Mudskippers breathe by absorbing oxygen across the skin (similar to frogs). A number of fishes have evolved so-called accessory breathing organs that extract oxygen from the air. Labyrinth fish (such as gouramis and bettas) have a labyrinth organ above the gills that performs this function. A few other fish have structures resembling labyrinth organs in form and function, most notably snakeheads, pikeheads, and the Clariidae catfish family.

Breathing air is primarily of use to fish that inhabit shallow, seasonally variable waters where the water's oxygen concentration may seasonally decline. Fishes dependent solely on dissolved oxygen, such as perch and cichlids, quickly suffocate, while air-breathers survive for much longer, in some cases in water that is little more than wet mud. At the most extreme, some air-breathing fish are able to survive in damp burrows for weeks without water, entering a state of aestivation (summertime hibernation) until water returns.

Tuna gills inside of the head. The fish head is oriented snout-downwards, with the view looking towards the mouth.

Fish can be divided into obligate air breathers and facultative air breathers. Obligate air breathers, such as the African lungfish, must breathe air periodically or they suffocate. Facultative air breathers, such as the catfish Hypostomus plecostomus, only breathe air if they need to and will 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.[12]

Circulation

Fish have a closed-loop circulatory system. The heart pumps the blood in a single loop throughout the body. In most fish, the heart consists of four parts, including two chambers and an entrance and exit.[13] The first part is the sinus venosus, a thin-walled sac that collects blood from the fish's veins before allowing it to flow to the second part, the atrium, which is a large muscular chamber. The atrium serves as a one-way antechamber, sends blood to the third part, ventricle. The ventricle is another thick-walled, muscular chamber and it pumps the blood, first to the fourth part, bulbous arteriosus, a large tube, and then out of the heart. The bulbus arteriosus connects to the aorta, through which blood flows to the gills for oxygenation.

Digestion

Jaws allow fish to eat a wide variety of food, including plants and other organisms. Fish ingest food through the mouth and break it down in the esophagus. In the stomach, food is further digested and, in many fish, processed in finger-shaped pouches called pyloric caeca, which secrete digestive enzymes and absorb nutrients. Organs such as the liver and pancreas add enzymes and various chemicals as the food moves through the digestive tract. The intestine completes the process of digestion and nutrient absorption.

Excretion

As with many aquatic animals, most fish release their nitrogenous wastes as ammonia. Some of the wastes diffuse through the gills. Blood wastes are filtered by the kidneys.

Saltwater fish tend to lose water because of osmosis. Their kidneys return water to the body. The reverse happens in freshwater fish: they tend to gain water osmotically. Their kidneys produce dilute urine for excretion. Some fish have specially adapted kidneys that vary in function, allowing them to move from freshwater to saltwater.

Scales

Main article: Scale (zoology)#Fish scales

The scales of fish originate from the mesoderm (skin); they may be similar in structure to teeth.

Sensory and nervous system

Dorsal view of the brain of the rainbow trout

Central nervous system

Fish typically have quite small brains relative to body size compared with other vertebrates, typically one-fifteenth the brain mass of a similarly sized bird or mammal.[14] However, some fish have relatively large brains, most notably mormyrids and sharks, which have brains about as massive relative to body weight as birds and marsupials.[15]

Fish brains divide into several regions. At the front are the olfactory lobes, a pair of structures that receive and process signals from the nostrils via the two olfactory nerves.[14] The olfactory lobes are very large in fishes that hunt primarily by smell, such as hagfish, sharks, and catfish. Behind the olfactory lobes is the two-lobed telencephalon, the structural equivalent to the cerebrum in higher vertebrates. In fishes the telencephalon is concerned mostly with olfaction.[14] Together these structures form the forebrain.

Connecting the forebrain to the midbrain is the diencephalon (in the diagram, this structure is below the optic lobes and consequently not visible). The diencephalon performs functions associated with hormones and homeostasis.[14] The pineal body lies just above the diencephalon. This structure detects light, maintains circadian rhythms, and controls color changes.[14]

The midbrain or mesencephalon contains the two optic lobes. These are very large in species that hunt by sight, such as rainbow trout and cichlids.[14]

The hindbrain or metencephalon is particularly involved in swimming and balance.[14] The cerebellum is a single-lobed structure that is typically the biggest part of the brain.[14] Hagfish and lampreys have relatively small cerebellae, while the mormyrid cerebellum is massive and apparently involved in their electrical sense.[14]

The brain stem or myelencephalon is the brain's posterior.[14] As well as controlling some muscles and body organs, in bony fish at least, the brain stem governs respiration and osmoregulation.[14]

Sense organs

Most fish possess highly developed sense organs. Nearly all daylight fish have color vision that is at least as good as a human's. Many fish also have chemoreceptors that are responsible for extraordinary senses of taste and smell. Although they have ears, many fish may not hear very well. Most fish have sensitive receptors that form the lateral line system, which detects gentle currents and vibrations, and senses the motion of nearby fish and prey.[16] Some fish, such as catfish and sharks, have organs that detect low-level electric current.[17] Other fish, like the electric eel, can produce electric current.

Fish orient themselves using landmarks and may use mental maps based on multiple landmarks or symbols. Fish behavior in mazes reveals that they possess spatial memory and visual discrimination.[18]

Capacity for pain

Further information: Pain in fish

Experiments done by William Tavolga provide evidence that fish have pain and fear responses. For instance, in Tavolga’s experiments, toadfish grunted when electrically shocked and over time they came to grunt at the mere sight of an electrode.[19]

In 2003, Scottish scientists at the University of Edinburgh and the Roslin Institute concluded that rainbow trout exhibit behaviors often associated with pain in other animals. Bee venom and acetic acid injected into the lips resulted in fish rocking their bodies and rubbing their lips along the sides and floors of their tanks, which the researchers concluded were attempts to relieve pain, similar to what mammals would do.[20][21][22] Neurons fired in a pattern resembling human neuronal patterns.[22]

Professor James D. Rose of the University of Wyoming claimed the study was flawed since it did not provide proof that fish possess ``conscious awareness, particularly a kind of awareness that is meaningfully like ours``.[23] Rose argues that since fish brains are so different from human brains, fish are probably not conscious in the manner humans are, so that reactions similar to human reactions to pain instead have other causes. Rose had published a study a year earlier arguing that fish cannot feel pain because their brains lack a neocortex.[24] However, animal behaviorist Temple Grandin argues that fish could still have consciousness without a neocortex because ``different species can use different brain structures and systems to handle the same functions.``[22]

Animal welfare advocates raise concerns about the possible suffering of fish caused by angling. Some countries, such as Germany have banned specific types of fishing, and the British RSPCA now formally prosecutes individuals who are cruel to fish.[25]

Muscular system

Main article: Fish locomotion

Most fish move by alternately contracting paired sets of muscles on either side of the backbone. These contractions form S-shaped curves that move down the body. As each curve reaches the back fin, backward force is applied to the water, and in conjunction with the fins, moves the fish forward. The fish's fins function like an airplane's flaps. Fins also increase the tail's surface area, increasing speed. The streamlined body of the fish decreases the amount of friction from the water. Since body tissue is denser than water, fish must compensate for the difference or they will sink. Many bony fishes have an internal organ called a swim bladder that adjusts their buoyancy through manipulation of gases.

Homeothermy

A 3-tonne (3.0 LT; 3.3 ST) great white shark off Isla Guadalupe

Although most fish are exclusively aquatic and ectothermic, there are exceptions to both cases.

Fish from multiple groups can live out of the water for extended time periods. Amphibious fish such as the mudskipper can live and move about on land for up to several days.

Certain species of fish maintain elevated body temperatures. Endothermic teleosts (bony fishes) are all in the suborder Scombroidei and include the billfishes, tunas, and one species of ``primitive`` mackerel (Gasterochisma melampus). All sharks in the family Lamnidae – shortfin mako, long fin mako, white, porbeagle, and salmon shark – are endothermic, and evidence suggests the trait exists in family Alopiidae (thresher sharks). The degree of endothermy varies from the billfish, which warm only their eyes and brain, to bluefin tuna and porbeagle sharks who maintain body temperatures elevated in excess of 20 °C above ambient water temperatures. See also gigantothermy. Endothermy, though metabolically costly, is thought to provide advantages such as increased muscle strength, higher rates of central nervous system processing, and higher rates of digestion.

Reproductive system

Further information: Spawn (biology)

Organs

Organs: 1. Liver, 2. Gas bladder, 3. Roe, 4. Pyloric caeca, 5. Stomach, 6. Intestine

Fish reproductive organs include testes and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused.[26] There may also be a range of secondary organs that increase reproductive fitness.

In terms of spermatogonia distribution, the structure of teleosts testes has two types: in the most common, spermatogonia occur all along the seminiferous tubules, while in Atherinomorph fishes they are confined to the distal portion of these structures. Fishes can present cystic or semi-cystic spermatogenesis in relation to the release phase of germ cells in cysts to the seminiferous tubules lumen.[26]

Fish ovaries may be of three types: gymnovarian, secondary gymnovarian or cystovarian. In the first type, the oocytes are released directly into the coelomic cavity and then enter the ostium, then through the oviduct and are eliminated. Secondary gymnovarian ovaries shed ova into the coelom from which they go directly into the oviduct. In the third type, the oocytes are conveyed to the exterior through the oviduct.[27] Gymnovaries are the primitive condition found in lungfish, sturgeon, and bowfin. Cystovaries characterize most teleosts, where the ovary lumen has continuity with the oviduct.[26] Secondary gymnovaries are found in salmonids and a few other teleosts.

Oogonia development in teleosts fish varies according to the group, and the determination of oogenesis dynamics allows the understanding of maturation and fertilization processes. Changes in the nucleus, ooplasm, and the surrounding layers characterize the oocyte maturation process.[26]

Postovulatory follicles are structures formed after oocyte release; they do not have endocrine function, present a wide irregular lumen, and are rapidly reabosrbed in a process involving the apoptosis of follicular cells. A degenerative process called follicular atresia reabsorbs vitellogenic oocytes not spawned. This process can also occur, but less frequently, in oocytes in other development stages.[26]

Some fish are hermaphrodites, having both testes and ovaries either at different phases in their life cycle or, as in hamlets, have them simultaneously.

Reproductive method

Over 97% of all known fishes are oviparous,[28] that is, the eggs develop outside the mother's body. Examples of oviparous fishes include salmon, goldfish, cichlids, tuna, and eels. In the majority of these species, fertilization takes place outside the mother's body, with the male and female fish shedding their gametes into the surrounding water. However, a few oviparous fishes practice internal fertilization, with the male using some sort of intromittent organ to deliver sperm into the genital opening of the female, most notably the oviparous sharks, such as the horn shark, and oviparous rays, such as skates. In these cases, the male is equipped with a pair of modified pelvic fins known as claspers.

Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of 1 millimetre (0.039 in).

Egg of lamprey

Egg of catshark (mermaids' purses)

Egg of bullhead shark

Egg of chimaera

An example of zooplankton

The newly-hatched young of oviparous fish are called larvae. They are usually poorly formed, carry a large yolk sac (for nourishment) and are very different in appearance from juvenile and adult specimens. The larval period in oviparous fish is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on zooplankton prey, a process which depends on typically inadequate zooplankton density, starving many larvae.

In ovoviviparous fish the eggs develop inside the mother's body after internal fertilization but receive little or no nourishment directly from the mother, depending instead on the yolk. Each embryo develops in its own egg. Familiar examples of ovoviviparous fishes include guppies, angel sharks, and coelacanths.

Some species of fish are viviparous. In such species the mother retains the eggs and nourishes the embryos. Typically, viviparous fishes have a structure analogous to the placenta seen in mammals connecting the mother's blood supply with the that of the embryo. Examples of viviparous fishes include the surf-perches, splitfins, and lemon shark. Some viviparous fishes exhibit oophagy, in which the developing embryos eat other eggs produced by the mother. This has been observed primarily among sharks, such as the shortfin mako and porbeagle, but is known for a few bony fish as well, such as the halfbeak Nomorhamphus ebrardtii.[29] Intrauterine cannibalism is an even more unusual mode of vivipary, in which the largest embryos eat weaker and smaller siblings. This behavior is also most commonly found among sharks, such as the grey nurse shark, but has also been reported for Nomorhamphus ebrardtii.[29]

Aquarists commonly refer to ovoviviparous and viviparous fishes as livebearers.

Immune system

Immune organs vary by type of fish.[30] In the jawless fish (lampreys and hagfishes), true lymphoid organs are absent. These fish rely on regions of lymphoid tissue within other organs to produce immune cells. For example, erythrocytes, macrophages and plasma cells are produced in the anterior kidney (or pronephros) and some areas of the gut (where granulocytes mature.) They resemble primitive bone marrow in hagfish. Cartilaginous fish (sharks and rays) have a more advanced immune system. They have three specialized organs that are unique to chondrichthyes; the epigonal organs (lymphoid tissue similar to mammalian bone) that surround the gonads, the