Fish physiology
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Fish physiology is the scientific study of how the component parts of
fish 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 li ...
function together in the living fish. It can be contrasted with
fish anatomy 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 co ...
, which is the study of the form or
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
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.


Respiration

Most fish exchange gases using
gill 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 ...
s on either side of the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
(throat). Gills are tissues which consist of threadlike structures called filaments. These filaments have many functions and "are involved in ion and water transfer as well as oxygen, carbon dioxide, acid and ammonia exchange. Each filament contains a
capillary A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
network that provides a large
surface area The surface area of a solid object is a measure of the total area that the surface of the object occupies. The mathematical definition of surface area in the presence of curved surfaces is considerably more involved than the definition of arc ...
for exchanging
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
and
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
. Fish exchange gases by pulling oxygen-rich water through their mouths and pumping it over their gills. In some fish, capillary blood flows in the opposite direction to the water, causing
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 ...
. The gills push the oxygen-poor water out through openings in the sides of the pharynx. Fish from multiple groups can live out of the water for extended time periods.
Amphibious fish 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 ...
such as the
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 ...
can live and move about on land for up to several days, or live in stagnant or otherwise oxygen depleted water. Many such fish can breathe air via a variety of mechanisms. The skin of anguillid eels may absorb oxygen directly. The
buccal cavity The buccal space (also termed the buccinator space) is a fascial space of the head and neck (sometimes also termed fascial tissue spaces or tissue spaces). It is a potential space in the cheek, and is paired on each side. The buccal space is super ...
of the
electric eel The electric eels are a genus, ''Electrophorus'', of neotropical freshwater fish from South America in the family Gymnotidae. They are known for their ability to stun their prey by generating electricity, delivering shocks at up to 860 volt ...
may breathe air. Catfish of the families
Loricariidae The Loricariidae is the largest family of catfish (order Siluriformes), with 92 genera and just over 680 species. Loricariids originate from freshwater habitats of Costa Rica, Panama, and tropical and subtropical South America. These fish are not ...
,
Callichthyidae Callichthyidae is a family of catfishes (order Siluriformes), called armored catfishes due to the two rows of bony plates (or scutes) along the lengths of their bodies. It contains some of the most popular freshwater aquarium fish, such as many ...
, and
Scoloplacidae ''Scoloplax'' is the only genus in the catfish (order Siluriformes) family Scoloplacidae, the spiny dwarf catfishes. Species The six currently recognized species in this genus are: * ''Scoloplax baileyi'' Rocha, Lazzarotto & Rapp Py-Daniel, ...
absorb air through their digestive tracts.
Lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
, with the exception of the Australian lungfish, and
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
s have paired lungs similar to those of
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s and must surface to gulp fresh air through the mouth and pass spent air out through the gills.
Gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
and
bowfin The bowfin (''Amia calva'') is a bony fish, native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being the sole surviving species of the Halecomorphi ...
have a vascularized swim bladder that functions in the same way.
Loaches Loaches are fish of the superfamily Cobitoidea. They are freshwater, benthic (bottom-dwelling) fish found in rivers and creeks throughout Eurasia and northern Africa. Loaches are among the most diverse groups of fish; the 1249 known species of ...
, trahiras, and many
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
breathe by passing air through the gut. Mudskippers breathe by absorbing oxygen across the skin (similar to frogs). A number of fish have evolved so-called accessory breathing organs that extract oxygen from the air. Labyrinth fish (such as
gourami Gouramis, or gouramies , are a group of freshwater anabantiform fishes that comprise the family Osphronemidae. The fish are native to Asia—from the Indian Subcontinent to Southeast Asia and northeasterly towards Korea. The name "gourami", of ...
s and
betta ''Betta'' is a large genus of small, active, often colorful, freshwater ray-finned fishes, in the gourami family (biology), family (Osphronemidae). The best known ''Betta'' species is ''B. splendens,'' commonly known as the Siamese fighting fi ...
s) have a
labyrinth organ The Anabantoidei are a suborder of anabantiform ray-finned freshwater fish distinguished by their possession of a lung-like labyrinth organ, which enables them to breathe air. The fish in the Anabantoidei suborder are known as anabantoids or lab ...
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. Fish dependent solely on dissolved oxygen, such as perch and
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
s, 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 Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and ...
(summertime hibernation) until water returns. Air breathing fish can be divided into ''obligate'' air breathers and ''facultative'' air breathers. Obligate air breathers, such as the
African lungfish ''Protopterus'' is the genus of four species of lungfish found in Africa. ''Protopterus'' was formerly thought to be the sole genus in the family Protopteridae, but more recent studies have classified it with ''Lepidosiren'' in the family Lepid ...
, are obligated to breathe air periodically or they suffocate. Facultative air breathers, such as the catfish ''
Hypostomus plecostomus ''Hypostomus plecostomus'', also known as the suckermouth catfish or the common pleco, is a tropical freshwater fish belonging to the armored catfish family (Loricariidae), named for the longitudinal rows of armor-like scutes that cover the upp ...
'', 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. All basal vertebrates breathe with
gills 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 ...
. The gills are carried right behind the head, bordering the posterior margins of a series of openings from the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
to the exterior. Each gill is supported by a cartilaginous or bony
gill arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arc ...
. The gills of
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s typically develop in the walls of the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
, along a series of gill slits opening to the exterior. Most species employ a
countercurrent exchange Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each ...
system to enhance the diffusion of substances in and out of the gill, with blood and water flowing in opposite directions to each other, which increases the efficiency of oxygen-uptake from the water. Fresh oxygenated water taken in through the mouth is uninterruptedly "pumped" through the gills in one direction, while the blood in the lamellae flows in the opposite direction, creating the countercurrent blood and water flow, on which the fish's survival depends. The gills are composed of comb-like filaments, the
gill lamellae Lamellae on a gecko's foot. 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 ...
, which help increase their surface area for oxygen exchange. When a fish breathes, it draws in a mouthful of water at regular intervals. Then it draws the sides of its throat together, forcing the water through the gill openings, so that it passes over the gills to the outside. The
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
have three pairs of arches,
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...
have five to seven pairs, while the primitive
jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostomes ...
have seven. The vertebrate ancestor no doubt had more arches, as some of their
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
relatives have more than 50 pairs of gills.
Higher vertebrates Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
do not develop gills, the gill arches form during
fetal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
, and lay the basis of essential structures such as
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
s, the
thyroid gland The thyroid, or thyroid gland, is an endocrine gland in vertebrates. In humans it is in the neck and consists of two connected lobe (anatomy), lobes. The lower two thirds of the lobes are connected by a thin band of Connective tissue, tissue cal ...
, the
larynx The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
, the ''columella'' (corresponding to the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
in
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
) and in mammals the malleus and incus. Fish gill slits may be the evolutionary ancestors of the
tonsil The tonsils are a set of lymphoid organs facing into the aerodigestive tract, which is known as Waldeyer's tonsillar ring and consists of the adenoid tonsil, two tubal tonsils, two palatine tonsils, and the lingual tonsils. These organs play an ...
s,
thymus gland The thymus is a specialized primary lymphoid organ of the immune system. Within the thymus, thymus cell lymphocytes or ''T cells'' mature. T cells are critical to the adaptive immune system, where the body adapts to specific foreign invaders. T ...
, and
Eustachian tube In anatomy, the Eustachian tube, also known as the auditory tube or pharyngotympanic tube, is a tube that links the nasopharynx to the middle ear, of which it is also a part. In adult humans, the Eustachian tube is approximately long and in d ...
s, as well as many other structures derived from the embryonic
branchial pouch In the embryonic development of vertebrates, pharyngeal pouches form on the endodermal side between the pharyngeal arches. The pharyngeal grooves (or clefts) form the lateral ectodermal surface of the neck region to separate the arches. The pou ...
es. Scientists have investigated what part of the body is responsible for maintaining the respiratory rhythm. They found that
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. N ...
s located in the
brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...
of fish are responsible for the genesis of the respiratory rhythm. The position of these neurons is slightly different from the centers of respiratory genesis in mammals but they are located in the same brain compartment, which has caused debates about the
homology Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor * Sequence homology, biological homology between DNA, RNA, or protein sequences *Homologous chrom ...
of respiratory centers between aquatic and terrestrial species. In both aquatic and terrestrial respiration, the exact mechanisms by which neurons can generate this involuntary rhythm are still not completely understood (see Involuntary control of respiration). Another important feature of the respiratory rhythm is that it is modulated to adapt to the oxygen consumption of the body. As observed in mammals, fish "breathe" faster and heavier when they do
physical exercise Exercise is a body activity that enhances or maintains physical fitness and overall health and wellness. It is performed for various reasons, to aid growth and improve strength, develop muscles and the cardiovascular system, hone athletic ...
. The mechanisms by which these changes occur have been strongly debated over more than 100 years between scientists. The authors can be classified in 2 schools: #Those who think that the major part of the respiratory changes are pre-programmed in the brain, which would imply that neurons from
locomotion Locomotion means the act or ability of something to transport or move itself from place to place. Locomotion may refer to: Motion * Motion (physics) * Robot locomotion, of man-made devices By environment * Aquatic locomotion * Flight * Locomo ...
centers of the brain connect to
respiratory center The respiratory center is located in the medulla oblongata and pons, in the brainstem. The respiratory center is made up of three major respiratory groups of neurons, two in the medulla and one in the pons. In the medulla they are the dorsal ...
s in anticipation of movements. #Those who think that the major part of the respiratory changes result from the detection of muscle contraction, and that respiration is adapted as a consequence of muscular contraction and oxygen consumption. This would imply that the brain possesses some kind of detection mechanisms that would trigger a respiratory response when muscular contraction occurs. Many now agree that both mechanisms are probably present and complementary, or working alongside a mechanism that can detect changes in oxygen and/or carbon dioxide blood saturation.


Bony fish

In
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
, the gills lie in a branchial chamber covered by a bony operculum. The great majority of bony fish species have five pairs of gills, although a few have lost some over the course of evolution. The operculum can be important in adjusting the pressure of water inside of the pharynx to allow proper ventilation of the gills, so 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. 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. Marine
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s also use gills to excrete
electrolyte An electrolyte is a medium containing ions that is electrically conducting through the movement of those ions, but not conducting electrons. This includes most soluble salts, acids, and bases dissolved in a polar solvent, such as water. Upon dis ...
s. The gills' large surface area tends to create a problem for fish that seek to regulate the
osmolarity Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). The osmolarity of a solution is usually expressed as Osm/L ...
of their internal fluids. Saltwater is less dilute than these internal fluids, so saltwater fish lose large quantities of water osmotically through their gills. To regain the water, they drink large amounts of
seawater Seawater, or salt water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has appro ...
and excrete the
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
. Freshwater is more dilute than the internal fluids of fish, however, so freshwater fish gain water osmotically through their gills. In some primitive bony fishes and
amphibians 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, arbore ...
, the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e bear external gills, branching off from the gill arches. These are reduced in adulthood, their function taken over by the gills proper in fishes and by
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
s 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
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s.Clack, J. A. (2002): Gaining ground: the origin and evolution of tetrapods. ''Indiana University Press'', Bloomington, Indiana. 369 pp


Cartilaginous fish

Like other fish, sharks extract
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
from seawater as it passes over their
gill 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 ...
s. Unlike other fish, shark gill slits are not covered, but lie in a row behind the head. A modified slit called a
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
lies just behind the eye, which assists the shark with taking in water during
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
and plays a major role in bottom–dwelling sharks. Spiracles are reduced or missing in active
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
sharks. While the shark is moving, water passes through the mouth and over the gills in a process known as "ram ventilation". While at rest, most sharks pump water over their gills to ensure a constant supply of oxygenated water. A small number of species have lost the ability to pump water through their gills and must swim without rest. These species are ''obligate ram ventilators'' and would presumably
asphyxiate Asphyxia or asphyxiation is a condition of deficient supply of oxygen to the body which arises from abnormal breathing. Asphyxia causes generalized hypoxia, which affects primarily the tissues and organs. There are many circumstances that can i ...
if unable to move. Obligate ram ventilation is also true of some pelagic bony fish species. The respiration and circulation process begins when deoxygenated
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the c ...
travels to the shark's two-chambered
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
. Here the shark pumps blood to its gills via the ventral
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes ...
artery An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pul ...
where it branches into afferent brachial arteries. Reoxygenation takes place in the gills and the reoxygenated blood flows into the efferent brachial arteries, which come together to form the
dorsal aorta The dorsal aortae are paired (left and right) embryological vessels which progress to form the descending aorta. The paired dorsal aortae arise from aortic arches that in turn arise from the aortic sac. The primary dorsal aorta is located deep t ...
. The blood flows from the dorsal aorta throughout the body. The deoxygenated blood from the body then flows through the
posterior cardinal vein The postcardinal veins or posterior cardinal veins join with the corresponding right and left cardinal veins to form the left common cardinal veins, which empty in the sinus venosus. In the development of a human embryo, most of the posterior card ...
s and enters the posterior cardinal
sinuses Paranasal sinuses are a group of four paired air-filled spaces that surround the nasal cavity. The maxillary sinuses are located under the eyes; the frontal sinuses are above the eyes; the ethmoidal sinuses are between the eyes and the sphenoid ...
. From there blood enters the heart ventricle and the cycle repeats. ----
Shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s and
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six or seven pairs. Adjacent slits are separated by a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...
gill arch from which projects a long sheet-like
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatri ...
, 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 raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of the ...
s, small projecting elements that help to filter food from the water. A smaller opening, the
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
, lies in the back of the first
gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
. This bears a small ''
pseudobranch The pseudobranch, also pseudobranchia is the reduced first gill arch of a fish (on the inner surface of the opercle, near the junction of the preopercle) as well as a reduced "false" gill in some gastropods. In teleost fish, the pseudobranchs are m ...
'' that resembles a gill in structure, but only receives blood already oxygenated by the true gills. The spiracle is thought to be homologous to the ear opening in higher vertebrates. Most sharks rely on ram ventilation, forcing water into the mouth and over the gills by rapidly swimming forward. In slow-moving or bottom dwelling species, especially among skates and rays, the spiracle may be enlarged, and the fish breathes by sucking water through this opening, instead of through the mouth.
Chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
s differ from other cartilagenous fish, having lost both the spiracle and the fifth gill slit. The remaining slits are covered by an operculum, developed from the septum of the gill arch in front of the first gill.


Lampreys and hagfish

Lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s and
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
do not have gill slits as such. Instead, the gills are contained in spherical pouches, with a circular opening to the outside. Like the gill slits of higher fish, each pouch contains two gills. In some cases, the openings may be fused together, effectively forming an operculum. Lampreys have seven pairs of pouches, while hagfishes may have six to fourteen, depending on the species. In the hagfish, the pouches connect with the pharynx internally. 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.


Circulation

The circulatory systems of all
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s are ''closed'', just as in humans. Still, the systems of
fish 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 li ...
,
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s show various stages of the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of the
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
. In fish, the system has only one circuit, with the blood being pumped through the capillaries of the
gill 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 ...
s and on to the capillaries of the body tissues. This is known as ''single cycle'' circulation. The heart of fish is therefore only a single pump (consisting of two chambers). Fish have a closed-loop circulatory system. The
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
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. The first part is the
sinus venosus The sinus venosus is a large quadrangular cavity which precedes the atrium on the venous side of the chordate heart. In mammals, it exists distinctly only in the embryonic heart, where it is found between the two venae cavae. However, the sinus v ...
, a thin-walled sac that collects blood from the fish's
vein Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated b ...
s before allowing it to flow to the second part, the
atrium Atrium may refer to: Anatomy * Atrium (heart), an anatomical structure of the heart * Atrium, the genital structure next to the genital aperture in the reproductive system of gastropods * Atrium of the ventricular system of the brain * Pulmona ...
, 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,
bulbus arteriosus In the circulatory system of fish, the bulbus arteriosus is a pear shaped chamber that functions as a capacitor, maintaining continuous blood flow into the gill arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, ...
, a large tube, and then out of the heart. The bulbus arteriosus connects to the
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes ...
, through which blood flows to the
gills 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 ...
for oxygenation. In amphibians and most reptiles, a double circulatory system is used, but the heart is not always completely separated into two pumps. Amphibians have a three-chambered
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
.


Digestion

Jaws Jaws or Jaw may refer to: Anatomy * Jaw, an opposable articulated structure at the entrance of the mouth ** Mandible, the lower jaw Arts, entertainment, and media * Jaws (James Bond), a character in ''The Spy Who Loved Me'' and ''Moonraker'' * ...
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 The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
. In the stomach, food is further digested and, in many fish, processed in finger-shaped pouches called
pyloric caeca Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish a ...
, which secrete digestive
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
and absorb nutrients. Organs such as the
liver The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
and
pancreas The pancreas is an organ of the digestive system and endocrine system of vertebrates. In humans, it is located in the abdomen behind the stomach and functions as a gland. The pancreas is a mixed or heterocrine gland, i.e. it has both an end ...
add enzymes and various chemicals as the food moves through the digestive tract. The intestine completes the process of digestion and nutrient absorption. In most vertebrates, digestion is a four-stage process involving the main structures of the
digestive tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans a ...
, starting with ingestion, placing food into the mouth, and concluding with the excretion of undigested material through the anus. From the mouth, the food moves to the stomach, where as bolus it is broken down chemically. It then moves to the intestine, where the process of breaking the food down into simple molecules continues and the results are absorbed as nutrients into the circulatory and lymphatic system. Although the precise shape and size of the stomach varies widely among different vertebrates, the relative positions of the oesophageal and duodenal openings remain relatively constant. As a result, the organ always curves somewhat to the left before curving back to meet the pyloric sphincter. However,
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s,
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
es,
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
s,
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
es, and some
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
fish have no stomach at all, with the oesophagus opening directly into the intestine. These animals all consume diets that either require little storage of food, or no pre-digestion with gastric juices, or both. The
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the p ...
is the part of the digestive tract following the stomach and followed by the
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces before being r ...
, and is where much of the
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intest ...
and absorption of food takes place. In fish, the divisions of the small intestine are not clear, and the terms ''anterior'' or ''proximal'' intestine may be used instead of
duodenum The duodenum is the first section of the small intestine in most higher vertebrates, including mammals, reptiles, and birds. In fish, the divisions of the small intestine are not as clear, and the terms anterior intestine or proximal intestine m ...
. The small intestine is found in all
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s, although its form and length vary enormously between species. In teleosts, it is relatively short, typically around one and a half times the length of the fish's body. It commonly has a number of ''pyloric caeca'', small pouch-like structures along its length that help to increase the overall surface area of the organ for digesting food. There is no ileocaecal valve in teleosts, with the boundary between the small intestine and the
rectum The rectum is the final straight portion of the large intestine in humans and some other mammals, and the Gastrointestinal tract, gut in others. The adult human rectum is about long, and begins at the rectosigmoid junction (the end of the s ...
being marked only by the end of the digestive epithelium. There is no small intestine as such in non-teleost fish, such as
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s,
sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...
s, and
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
. Instead, the digestive part of the gut forms a spiral intestine, connecting the stomach to the rectum. In this type of gut, the intestine itself is relatively straight, but has a long fold running along the inner surface in a spiral fashion, sometimes for dozens of turns. This valve greatly increases both the surface area and the effective length of the intestine. The lining of the spiral intestine is similar to that of the small intestine in teleosts and non-mammalian tetrapods. In
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s, the spiral valve is extremely small, possibly because their diet requires little digestion.
Hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
have no spiral valve at all, with digestion occurring for almost the entire length of the intestine, which is not subdivided into different regions. The
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces before being r ...
is the last part of the
digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
normally found in vertebrate animals. Its function is to absorb water from the remaining indigestible food matter, and then to pass useless waste material from the body. In fish, there is no true large intestine, but simply a short rectum connecting the end of the digestive part of the gut to the cloaca. In
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s, this includes a ''rectal gland'' that secretes salt to help the animal maintain
osmotic Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
balance with the seawater. The gland somewhat resembles a caecum in structure, but is not a homologous structure. As with many aquatic animals, most fish release their nitrogenous wastes as
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
. Some of the wastes
diffuse Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
through the gills. Blood wastes are
filtered Filtration is a physical separation process that separates solid matter and fluid from a mixture using a ''filter medium'' that has a complex structure through which only the fluid can pass. Solid particles that cannot pass through the filter m ...
by the
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
s. Saltwater fish tend to lose water because of
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
. Their kidneys return water to the body. The reverse happens in
freshwater fish Freshwater fish are those that spend some or all of their lives in fresh water, such as rivers and lakes, with a salinity of less than 1.05%. These environments differ from marine conditions in many ways, especially the difference in levels of s ...
: 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. In sharks, digestion can take a long time. The food moves from the mouth to a J-shaped stomach, where it is stored and initial digestion occurs. Unwanted items may never get past the stomach, and instead the shark either vomits or turns its stomachs inside out and ejects unwanted items from its mouth. One of the biggest differences between the digestive systems of sharks and mammals is that sharks have much shorter intestines. This short length is achieved by the
spiral valve A spiral valve or scroll valve is the corkscrew-shaped lower portion of the intestine of some sharks, Acipenseriformes (sturgeon and paddlefish), rays, skates, bichirs, Lepisosteiformes (gars), and lungfishes. A modification of the ileum, the s ...
with multiple turns within a single short section instead of a long tube-like intestine. The valve provides a long surface area, requiring food to circulate inside the short gut until fully digested, when remaining waste products pass into the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
.


Endocrine system


Regulation of social behaviour

Oxytocin Oxytocin (Oxt or OT) is a peptide hormone and neuropeptide normally produced in the hypothalamus and released by the posterior pituitary. It plays a role in social bonding, reproduction, childbirth, and the period after childbirth. Oxytocin ...
is a group of
neuropeptide Neuropeptides are chemical messengers made up of small chains of amino acids that are synthesized and released by neurons. Neuropeptides typically bind to G protein-coupled receptors (GPCRs) to modulate neural activity and other tissues like the ...
s found in most vertebrates. One form of oxytocin functions as a
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
which is associated with human love. In 2012, researchers injected
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
s from the social species ''
Neolamprologus pulcher ''Neolamprologus pulcher'' is a species of cichlid endemic to Lake Tanganyika where it prefers locations with plenty of sedimentation. The common name for ''N. pulcher'' is the daffodil cichlid. This species can reach a length of TL. It can ...
'', either with this form of isotocin or with a control saline solution. They found isotocin increased "responsiveness to social information", which suggests "it is a key regulator of social behavior that has evolved and endured since ancient times".


Effects of pollution

Fish can
bioaccumulate Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance at a rate faster than that at which the substance is lost or eliminated ...
pollutants that are discharged into waterways. Estrogenic compounds found in pesticides, birth control, plastics, plants, fungi, bacteria, and synthetic drugs leeched into rivers are affecting the
endocrine system The endocrine system is a messenger system comprising feedback loops of the hormones released by internal glands of an organism directly into the circulatory system, regulating distant target organs. In vertebrates, the hypothalamus is the neu ...
s of native species. In Boulder, Colorado,
white sucker The white sucker (''Catostomus commersonii)'' is a species of freshwater cypriniform fish inhabiting the upper Midwest and Northeast in North America, but it is also found as far south as Georgia and as far west as New Mexico. The fish is commonl ...
fish found downstream of a municipal waste water treatment plant exhibit impaired or abnormal sexual development. The fish have been exposed to higher levels of estrogen, and leading to feminized fish. Males display female reproductive organs, and both sexes have reduced fertility, and a higher hatch mortality. Freshwater habitats in the United States are widely contaminated by the common pesticide
atrazine Atrazine is a chlorinated herbicide of the triazine class. It is used to prevent pre-emergence broadleaf weeds in crops such as maize (corn), soybean and sugarcane and on turf, such as golf courses and residential lawns. Atrazine's primary manufa ...
. There is controversy over the degree to which this pesticide harms the endocrine systems of freshwater fish and
amphibians 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, arbore ...
. Non-industry-funded researchers consistently report harmful effects while industry-funded researchers consistently report no harmful effects. In the marine ecosystem,
organochlorine An organochloride, organochlorine compound, chlorocarbon, or chlorinated hydrocarbon is an organic compound containing at least one covalent bond, covalently bonded atom of chlorine. The chloroalkane class (alkanes with one or more hydrogens subst ...
contaminants like pesticides, herbicides (DDT), and
chlordan Chlordane, or chlordan, is an organochlorine compound that was used as a pesticide. It is a white solid. In the United States, chlordane was used for termite-treatment of approximately 30 million homes until it was banned in 1988. Chlordane was b ...
are accumulating within fish tissue and disrupting their endocrine system. High frequencies of infertility and high levels of organochlorines have been found in
bonnethead shark The bonnethead (''Sphyrna tiburo''), also called a bonnet shark or shovelhead, is a small member of the hammerhead shark genus ''Sphyrna'', and part of the family Sphyrnidae. It is an abundant species in the littoral zone of the North Atlantic ...
s along the Gulf Coast of Florida. These endocrine-disrupting compounds are similar in structure to naturally occurring hormones in fish. They can modulate hormonal interactions in fish by: * binding to cellular receptors, causing unpredictable and abnormal cell activity * blocking receptor sites, inhibiting activity * promoting the creation of extra receptor sites, amplifying the effects of the hormone or compound * interacting with naturally occurring hormones, changing their shape and impact * affecting hormone synthesis or metabolism, causing an improper balance or quantity of hormones


Osmoregulation

Two major types of
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
are osmoconformers and osmoregulators.
Osmoconformer Osmoconformers are marine organisms that maintain an internal environment which is isotonic to their external environment. This means that the osmotic pressure of the organism's cells is equal to the osmotic pressure of their surrounding environme ...
s match their body osmolarity to their environment actively or passively. Most marine invertebrates are osmoconformers, although their ionic composition may be different from that of seawater.
Osmoregulator Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration of ...
s tightly regulate their body
osmolarity Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). The osmolarity of a solution is usually expressed as Osm/L ...
, which always stays constant, and are more common in the animal kingdom. Osmoregulators actively control salt concentrations despite the salt concentrations in the environment. An example is freshwater fish. The gills actively uptake salt from the environment by the use of mitochondria-rich cells. Water will diffuse into the fish, so it excretes a very
hypotonic In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane-imp ...
(dilute) urine to expel all the excess water. A marine
fish 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 li ...
has an internal osmotic concentration lower than that of the surrounding seawater, so it tends to lose water and gain salt. It actively excretes
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
out from the
gill 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 ...
s. Most fish are stenohaline, which means they are restricted to either salt or fresh water and cannot survive in water with a different salt concentration than they are adapted to. However, some fish show a tremendous ability to effectively osmoregulate across a broad range of salinities; fish with this ability are known as
euryhaline Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the molly (''Poecilia sphenops'') which can live in fresh water, brackish water, or salt water. The green crab (''Carcinus maenas'') is an e ...
species, e.g.,
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
. Salmon has been observed to inhabit two utterly disparate environments — marine and fresh water — and it is inherent to adapt to both by bringing in behavioral and physiological modifications. In contrast to bony fish, with the exception of the
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
, the blood and other tissue of sharks and
Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
is generally isotonic to their marine environments because of the high concentration of
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important r ...
and
trimethylamine Trimethylamine (TMA) is an organic compound with the formula N(CH3)3. It is a colorless, hygroscopic, and flammable tertiary amine. It is a gas at room temperature but is usually sold as a 40% solution in water. (It is also sold in pressurized ...
N-oxide (TMAO), allowing them to be in
osmotic Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
balance with the seawater. This adaptation prevents most sharks from surviving in freshwater, and they are therefore confined to
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
environments. A few exceptions exist, such as the
bull shark The bull shark (''Carcharhinus leucas''), also known as the Zambezi shark (informally zambi) in Africa and Lake Nicaragua shark in Nicaragua, is a species of requiem shark commonly found worldwide in warm, shallow waters along coasts and in riv ...
, which has developed a way to change its
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
function to excrete large amounts of urea. When a shark dies, the urea is broken down to ammonia by bacteria, causing the dead body to gradually smell strongly of ammonia. Sharks have adopted a different, efficient mechanism to conserve water, i.e., osmoregulation. They retain urea in their blood in relatively higher concentration. Urea is damaging to living tissue so, to cope with this problem, some fish retain ''
trimethylamine oxide Trimethylamine ''N''-oxide (TMAO) is an organic compound with the formula (CH3)3NO. It is in the class of amine oxides. Although the anhydrous compound is known, trimethylamine ''N''-oxide is usually encountered as the dihydrate. Both the anhydro ...
''. This provides a better solution to urea's toxicity. Sharks, having slightly higher solute concentration (i.e., above 1000 mOsm which is sea solute concentration), do not drink water like fresh water fish.


Thermoregulation

Homeothermy Homeothermy, homothermy or homoiothermy is thermoregulation that maintains a stable internal body temperature regardless of external influence. This internal body temperature is often, though not necessarily, higher than the immediate environmen ...
and
poikilothermy A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
refer to how stable an organism's temperature is. Most
endothermic In thermochemistry, an endothermic process () is any thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, H.P., Butler, L. J. (2015).''Principle of Modern Chemistry'', Brooks Cole. p. ...
organisms are homeothermic, like
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. However, animals with facultative
endothermy An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
are often poikilothermic, meaning their temperature can vary considerably. Similarly, most fish are
ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
s, as all of their heat comes from the surrounding water. However, most are homeotherms because their temperature is very stable. Most organisms have a preferred temperature range, however some can be acclimated to temperatures colder or warmer than what they are typically used to. An organism's preferred temperature is typically the temperature at which the organism's physiological processes can act at optimal rates. When fish become acclimated to other temperatures, the efficiency of their physiological processes may decrease but will continue to function. This is called the
thermal neutral zone Endothermic organisms known as homeotherms maintain internal temperatures with minimal metabolic regulation within a range of ambient temperatures called the thermal neutral zone (TNZ). Within the TNZ the basal rate of heat production is equal to ...
at which an organism can survive indefinitely. H.M. Vernon has done work on the death temperature and paralysis temperature (temperature of heat rigor) of various animals. He found that species of the same
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
showed very similar temperature values, those from the
Amphibia 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, arbore ...
examined being 38.5 °C, fish 39 °C,
Reptilia Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
45 °C, and various
Mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s 46 °C. To cope with low temperatures, some
fish 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 li ...
have developed the ability to remain functional even when the water temperature is below freezing; some use natural
antifreeze An antifreeze is an additive which lowers the freezing point of a water-based liquid. An antifreeze mixture is used to achieve freezing-point depression for cold environments. Common antifreezes also increase the boiling point of the liquid, all ...
or
antifreeze proteins Antifreeze proteins (AFPs) or ice structuring proteins refer to a class of polypeptides produced by certain animals, plants, fungi and bacteria that permit their survival in temperatures below the freezing point of water. AFPs bind to small i ...
to resist ice crystal formation in their tissues. Most sharks are "cold-blooded" or, more precisely,
poikilotherm A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
ic, meaning that their internal
body temperature Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
matches that of their ambient environment. Members of the family
Lamnidae The Lamnidae are the family of mackerel sharks known as white sharks. They are large, fast-swimming predatory fish found in oceans worldwide, though prefer environments with colder water. The name of the family is formed from the Greek word ''lam ...
(such as the
shortfin mako shark The shortfin mako shark (; ; ''Isurus oxyrinchus''), also known as the blue pointer or bonito shark, is a large mackerel shark. It is commonly referred to as the mako shark, as is the longfin mako shark (''Isurus paucus''). The shortfin mako can ...
and the
great white shark The great white shark (''Carcharodon carcharias''), also known as the white shark, white pointer, or simply great white, is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is nota ...
) are
homeothermic Homeothermy, homothermy or homoiothermy is thermoregulation that maintains a stable internal body temperature regardless of external influence. This internal body temperature is often, though not necessarily, higher than the immediate environmen ...
and maintain a higher body temperature than the surrounding water. In these sharks, a strip of
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cellu ...
red muscle located near the center of the body generates the heat, which the body retains via a
countercurrent exchange Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each ...
mechanism by a system of
blood vessel The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away ...
s called the
rete mirabile A rete mirabile (Latin for "wonderful net"; plural retia mirabilia) is a complex of arteries and veins lying very close to each other, found in some vertebrates, mainly warm-blooded ones. The rete mirabile utilizes countercurrent blood flow within ...
("miraculous net"). The common thresher shark has a similar mechanism for maintaining an elevated body temperature, which is thought to have evolved independently.
Tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
can maintain the temperature of certain parts of their body above the temperature of ambient seawater. For example,
bluefin tuna Bluefin tuna is a common name used to refer to several species of tuna of the genus ''Thunnus ''Thunnus'' is a genus of ocean-dwelling, ray-finned bony fish from the mackerel family, Scombridae. More specifically, ''Thunnus'' is one of five ...
maintain a core body temperature of , in water as cold as . However, unlike typical endothermic creatures such as mammals and birds, tuna do not maintain temperature within a relatively narrow range. Tuna achieve
endothermy An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
by conserving the heat generated through normal
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
. The
rete mirabile A rete mirabile (Latin for "wonderful net"; plural retia mirabilia) is a complex of arteries and veins lying very close to each other, found in some vertebrates, mainly warm-blooded ones. The rete mirabile utilizes countercurrent blood flow within ...
("wonderful net"), the intertwining of veins and arteries in the body's periphery, transfers heat from
venous blood Venous blood is deoxygenated blood which travels from the peripheral blood vessels, through the venous system into the right atrium of the heart. Deoxygenated blood is then pumped by the right ventricle to the lungs via the pulmonary artery whic ...
to
arterial blood Arterial blood is the oxygenated blood in the circulatory system found in the pulmonary vein, the left chambers of the heart, and in the artery, arteries. It is bright red in color, while venous blood is dark red in color (but looks purple through ...
via a
counter-current exchange Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each ...
system, thus mitigating the effects of surface cooling. This allows the tuna to elevate the temperatures of the highly
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cellu ...
tissues of the skeletal muscles, eyes and brain, which supports faster swimming speeds and reduced energy expenditure, and which enables them to survive in cooler waters over a wider range of ocean environments than those of other fish. In all tunas, however, the heart operates at
ambient temperature Colloquially, "room temperature" is a range of air temperatures that most people prefer for indoor settings. It feels comfortable to a person when they are wearing typical indoor clothing. Human comfort can extend beyond this range depending on ...
, as it receives cooled blood, and coronary circulation is directly from the
gills 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 ...
. *
Homeothermy Homeothermy, homothermy or homoiothermy is thermoregulation that maintains a stable internal body temperature regardless of external influence. This internal body temperature is often, though not necessarily, higher than the immediate environmen ...
: Although most fish are exclusively
ectothermic An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
, there are exceptions. Certain species of fish maintain elevated body temperatures.
Endothermic In thermochemistry, an endothermic process () is any thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, H.P., Butler, L. J. (2015).''Principle of Modern Chemistry'', Brooks Cole. p. ...
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s (bony fish) are all in the suborder
Scombroidei Scombroidei is a suborder of the Perciformes, the largest order of fish. The suborder includes the barracuda, tuna, and mackerel, as well as the billfish The term billfish refers to a group of saltwater predatory fish characterised by pro ...
and include the
billfish The term billfish refers to a group of saltwater predatory fish characterised by prominent pointed bills (rostra), and by their large size; some are longer than . Extant billfish include sailfish and marlin, which make up the family Istiophor ...
es, tunas, including a "primitive"
mackerel Mackerel is a common name applied to a number of different species of pelagic fish, mostly from the family Scombridae. They are found in both temperate and tropical seas, mostly living along the coast or offshore in the oceanic environment. ...
species, '' Gasterochisma melampus''. All sharks in the family
Lamnidae The Lamnidae are the family of mackerel sharks known as white sharks. They are large, fast-swimming predatory fish found in oceans worldwide, though prefer environments with colder water. The name of the family is formed from the Greek word ''lam ...
– shortfin mako, long fin mako, white, porbeagle, and salmon shark – are endothermic, and evidence suggests the trait exists in family
Alopiidae Thresher sharks are large lamniform sharks of the family Alopiidae found in all temperate and tropical oceans of the world; the family contains three extant species, all within the genus ''Alopias''. All three thresher shark species have been l ...
(
thresher sharks Thresher sharks are large lamniform sharks of the family Alopiidae found in all temperate and tropical oceans of the world; the family contains three extant species, all within the genus ''Alopias''. All three thresher shark species have been ...
). The degree of endothermy varies from the
billfish The term billfish refers to a group of saltwater predatory fish characterised by prominent pointed bills (rostra), and by their large size; some are longer than . Extant billfish include sailfish and marlin, which make up the family Istiophor ...
, which warm only their eyes and brain, to
bluefin tuna Bluefin tuna is a common name used to refer to several species of tuna of the genus ''Thunnus ''Thunnus'' is a genus of ocean-dwelling, ray-finned bony fish from the mackerel family, Scombridae. More specifically, ''Thunnus'' is one of five ...
and
porbeagle shark The porbeagle (''Lamna nasus'') is a species of mackerel shark in the family Lamnidae, distributed widely in the cold and temperate marine waters of the North Atlantic and Southern Hemisphere. In the North Pacific, its ecological equivalent is ...
s who maintain body temperatures elevated in excess of above ambient water temperatures. ''See also
gigantothermy Gigantothermy (sometimes called ectothermic homeothermy or inertial homeothermy) is a phenomenon with significance in biology and paleontology, whereby large, bulky ectothermic animals are more easily able to maintain a constant, relatively high bod ...
''. Endothermy, though metabolically costly, is thought to provide advantages such as increased muscle strength, higher rates of central
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
processing, and higher rates of
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intest ...
. In some fish, a
rete mirabile A rete mirabile (Latin for "wonderful net"; plural retia mirabilia) is a complex of arteries and veins lying very close to each other, found in some vertebrates, mainly warm-blooded ones. The rete mirabile utilizes countercurrent blood flow within ...
allows for an increase in muscle temperature in regions where this network of vein and arteries is found. The fish is able to
thermoregulate Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
certain areas of their body. Additionally, this increase in temperature leads to an increase in basal metabolic temperature. The fish is now able to split ATP at a higher rate and ultimately can swim faster. The eye of a
swordfish Swordfish (''Xiphias gladius''), also known as broadbills in some countries, are large, highly migratory predatory fish characterized by a long, flat, pointed bill. They are a popular sport fish of the billfish category, though elusive. Swordfis ...
can generate
heat In thermodynamics, heat is defined as the form of energy crossing the boundary of a thermodynamic system by virtue of a temperature difference across the boundary. A thermodynamic system does not ''contain'' heat. Nevertheless, the term is al ...
to better cope with detecting their
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
at depths of .


Muscular system

Fish swim by contracting longitudinal red muscle and obliquely oriented white muscles. The red muscle is aerobic and needs oxygen which is supplied by
myoglobin Myoglobin (symbol Mb or MB) is an iron- and oxygen-binding protein found in the cardiac and skeletal muscle tissue of vertebrates in general and in almost all mammals. Myoglobin is distantly related to hemoglobin. Compared to hemoglobin, myoglobi ...
. The white muscle is anaerobic and it does not need oxygen. Red muscles are used for sustained activity such as cruising at slow speeds on ocean migrations. White muscles are used for bursts of activity, such as jumping or sudden bursts of speed for catching prey. Mostly fish have white muscles, but the muscles of some fishes, such as scombroids and
salmonid Salmonidae is a family of ray-finned fish that constitutes the only currently extant family in the order Salmoniformes . It includes salmon (both Atlantic and Pacific species), trout (both ocean-going and landlocked), chars, freshwater whitefis ...
s, range from pink to dark red. The red myotomal muscles derive their colour from
myoglobin Myoglobin (symbol Mb or MB) is an iron- and oxygen-binding protein found in the cardiac and skeletal muscle tissue of vertebrates in general and in almost all mammals. Myoglobin is distantly related to hemoglobin. Compared to hemoglobin, myoglobi ...
, an oxygen-binding molecule, which tuna express in quantities far higher than most other fish. The oxygen-rich blood further enables energy delivery to their muscles. 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. A typical characteristic of many animals that utilize undulatory locomotion is that they have segmented muscles, or blocks of
myomere Myomeres are blocks of skeletal muscle tissue arranged in sequence, commonly found in aquatic chordates. Myomeres are separated from adjacent myomeres by connective fascia (myosepta) and most easily seen in larval fishes or in the olm. Myomere ...
s, running from their head to tails which are separated by connective tissue called myosepta. In addition, some segmented muscle groups, such the lateral hypaxial musculature in the salamander are oriented at an angle to the longitudinal direction. For these obliquely oriented fibers the strain in the longitudinal direction is greater than the strain in the muscle fiber direction leading to an architectural gear ratio greater than 1. A higher initial angle of orientation and more dorsoventral bulging produces a faster muscle contraction but results in a lower amount of force production. It is hypothesized that animals employ a variable gearing mechanism that allows self-regulation of force and velocity to meet the mechanical demands of the contraction. When a
pennate muscle A pennate or pinnate muscle (also called a penniform muscle) is a type of skeletal muscle with fascicles that attach obliquely (in a slanting position) to its tendon. This type of muscle generally allows higher force production but a smaller ra ...
is subjected to a low force, resistance to width changes in the muscle cause it to rotate which consequently produce a higher architectural gear ratio (AGR) (high velocity). However, when subject to a high force, the perpendicular fiber force component overcomes the resistance to width changes and the muscle compresses producing a lower AGR (capable of maintaining a higher force output). Most fishes bend as a simple, homogenous beam during swimming via contractions of longitudinal red muscle fibers and obliquely oriented white muscle fibers within the segmented axial musculature. The fiber
strain Strain may refer to: Science and technology * Strain (biology), variants of plants, viruses or bacteria; or an inbred animal used for experimental purposes * Strain (chemistry), a chemical stress of a molecule * Strain (injury), an injury to a mu ...
(εf) experienced by the longitudinal red muscle fibers is equivalent to the longitudinal strain (εx). The deeper white muscle fibers fishes show diversity in arrangement. These fibers are organized into cone-shaped structures and attach to connective tissue sheets known as myosepta; each fiber shows a characteristic dorsoventral (α) and mediolateral (φ) trajectory. The segmented architecture theory predicts that, εx > εf. This phenomenon results in an architectural gear ratio, determined as longitudinal strain divided by fiber strain (εx / εf), greater than one and longitudinal velocity amplification; furthermore, this emergent velocity amplification may be augmented by variable architectural gearing via mesolateral and dorsoventral shape changes, a pattern seen in
pennate muscle A pennate or pinnate muscle (also called a penniform muscle) is a type of skeletal muscle with fascicles that attach obliquely (in a slanting position) to its tendon. This type of muscle generally allows higher force production but a smaller ra ...
contractions. A red-to-white gearing ratio (red εf / white εf) captures the combined effect of the longitudinal red muscle fiber and oblique white muscle fiber strains.


Buoyancy

The body of a fish is denser than water, so fish must compensate for the difference or they will sink. Many
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
es have an internal organ called a
swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled Organ (anatomy), organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their curren ...
, or gas bladder, that adjusts their buoyancy through manipulation of gases. In this way, fish can stay at the current water depth, or ascend or descend without having to waste energy in swimming. The bladder is only found in bony fishes. In the more primitive groups like some
minnows Minnow is the common name for a number of species of small freshwater fish, belonging to several genera of the families Cyprinidae and Leuciscidae. They are also known in Ireland as pinkeens. Smaller fish in the subfamily Leusciscidae are ...
,
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
s and
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
, the bladder is open to the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
and double as a
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
. It is often absent in fast swimming fishes such as the tuna and mackerel families. The condition of a bladder open to the esophagus is called
physostome Physostomes are fishes that have a pneumatic duct connecting the gas bladder to the alimentary canal. This allows the gas bladder to be filled or emptied via the mouth. This not only allows the fish to fill their bladder by gulping air, but also ...
, the closed condition
physoclist Physoclisti are, collectively, fishes that lack a connection between the gas bladder and the alimentary canal, with the bladder serving only as a buoyancy organ. Addition and removal of the gases from the gas bladder in such physoclistous fishes ...
. In the latter, the gas content of the bladder is controlled through the rete mirabilis, a network of blood vessels effecting gas exchange between the bladder and the blood. In some fish, a rete mirabile fills the swim bladder with oxygen. A countercurrent exchange system is utilized between the venous and arterial capillaries. By lowering the pH levels in the venous capillaries, oxygen unbinds from blood
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
. This causes an increase in venous blood oxygen concentration, allowing the oxygen to diffuse through the capillary membrane and into the arterial capillaries, where oxygen is still sequestered to hemoglobin. The cycle of diffusion continues until the concentration of oxygen in the arterial capillaries is
supersaturated In physical chemistry, supersaturation occurs with a solution when the concentration of a solute exceeds the concentration specified by the value of solubility at equilibrium. Most commonly the term is applied to a solution of a solid in a liqu ...
(larger than the concentration of oxygen in the swim bladder). At this point, the free oxygen in the arterial capillaries diffuses into the swim bladder via the gas gland. Unlike bony fish, sharks do not have gas-filled swim bladders for buoyancy. Instead, sharks rely on a large liver filled with oil that contains
squalene Squalene is an organic compound. It is a triterpenoid with the formula C30H50. It is a colourless oil, although impure samples appear yellow. It was originally obtained from shark liver oil (hence its name, as ''Squalus'' is a genus of sharks). A ...
, and their cartilage, which is about half the normal density of bone. Their liver constitutes up to 30% of their total body mass. The liver's effectiveness is limited, so sharks employ
dynamic lift A fluid flowing around an object exerts a force on it. Lift is the Vector (geometric)#Vector components, component of this force that is perpendicular to the oncoming flow direction. It contrasts with the drag (physics), drag force, which is th ...
to maintain depth when not swimming.
Sand tiger shark The sand tiger shark (''Carcharias taurus''), gray nurse shark, spotted ragged-tooth shark or blue-nurse sand tiger, is a species of shark that inhabits subtropical and temperate waters worldwide. It inhabits the continental shelf, from sandy sho ...
s store air in their stomachs, using it as a form of swim bladder. Most sharks need to constantly swim in order to breathe and cannot sleep very long without sinking (if at all). However, certain species, like the
nurse shark The nurse shark (''Ginglymostoma cirratum'') is an elasmobranch fish in the family Ginglymostomatidae. The conservation status of the nurse shark is globally assessed as Vulnerable in the IUCN List of Threatened Species. They are considered t ...
, are capable of pumping water across their gills, allowing them to rest on the ocean bottom.


Sensory systems

Most fish possess highly developed sense organs. Nearly all daylight fish have color vision that is at least as good as a human's (see
vision in fishes Vision is an important sensory system for most species of fish. Fish eyes are similar to the eyes of terrestrial vertebrates like birds and mammals, but have a more spherical lens. Birds and mammals (including humans) normally adjust focus by ...
). 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 The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
, which detects gentle currents and vibrations, and senses the motion of nearby fish and prey. Sharks can sense frequencies in the range of 25 to 50  Hz through their lateral line. 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.


Vision

Vision Vision, Visions, or The Vision may refer to: Perception Optical perception * Visual perception, the sense of sight * Visual system, the physical mechanism of eyesight * Computer vision, a field dealing with how computers can be made to gain un ...
is an important
sensory system The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons (including the sensory receptor cells), neural pathways, and parts of the brain involved i ...
for most species of fish. Fish eyes are similar to those of
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s like
birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
and mammals, but have a more
spherical A sphere () is a geometrical object that is a three-dimensional analogue to a two-dimensional circle. A sphere is the set of points that are all at the same distance from a given point in three-dimensional space.. That given point is the ce ...
lens A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
. Their
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
s generally have both
rod cell Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in per ...
s and
cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...
s (for
scotopic In the study of human visual perception, scotopic vision (or scotopia) is the vision of the eye under low-light conditions. The term comes from Greek ''skotos'', meaning "darkness", and ''-opia'', meaning "a condition of sight". In the human eye, ...
and
photopic vision Photopic vision is the vision of the eye under well-lit conditions (luminance levels from 10 to 108  cd/m2). In humans and many other animals, photopic vision allows color perception, mediated by cone cells, and a significantly higher visua ...
), and most species have
colour vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of ...
. Some fish can see
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
and some can see
polarized light Polarization (also polarisation) is a property applying to transverse waves that specifies the geometrical orientation of the oscillations. In a transverse wave, the direction of the oscillation is perpendicular to the direction of motion of the ...
. Amongst
jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostomes ...
, the
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
has well-developed eyes, while the
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
has only primitive eyespots. Fish vision shows
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
to their visual environment, for example
deep sea fish Deep-sea fish are fish that live in the darkness below the sunlit surface waters, that is below the epipelagic or photic zone of the sea. The lanternfish is, by far, the most common deep-sea fish. Other deep sea fishes include the flashlight ...
es have eyes suited to the dark environment.


Hearing

Hearing Hearing, or auditory perception, is the ability to perceive sounds In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psycholog ...
is an important sensory system for most species of fish. Hearing threshold and the ability to localize sound sources are reduced underwater, in which the speed of sound is faster than in air. Underwater hearing is by
bone conduction Bone conduction is the conduction of sound to the inner ear primarily through the bones of the skull, allowing the hearer to perceive audio content without blocking the ear canal. Bone conduction transmission occurs constantly as sound waves vibra ...
, and localization of sound appears to depend on differences in amplitude detected by bone conduction. Aquatic animals such as fish, however, have a more specialized hearing apparatus that is effective underwater. Fish can sense sound through their
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
s and their
otolith An otolith ( grc-gre, ὠτο-, ' ear + , ', a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The sa ...
s (ears). Some fishes, such as some species of
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
and
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Oceans, i ...
, hear through their swim bladders, which function rather like a hearing aid. Hearing is well-developed in
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
, which have the Weberian organ, three specialized vertebral processes that transfer vibrations in the swim bladder to the inner ear. Although it is hard to test sharks' hearing, they may have a sharp
sense of hearing A sense is a biological system used by an organism for sensation, the process of gathering information about the world through the detection of stimuli. (For example, in the human body, the brain which is part of the central nervous system rec ...
and can possibly hear prey many miles away. A small opening on each side of their heads (not the
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
) leads directly into the
inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
through a thin channel. The
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
shows a similar arrangement, and is open to the environment via a series of openings called lateral line pores. This is a reminder of the common origin of these two vibration- and sound-detecting organs that are grouped together as the acoustico-lateralis system. In bony fish and
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s the external opening into the inner ear has been lost.


Chemoreception

Sharks have keen
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
senses, located in the short duct (which is not fused, unlike bony fish) between the anterior and posterior nasal openings, with some species able to detect as little as one
part per million In science and engineering, the parts-per notation is a set of pseudo-units to describe small values of miscellaneous dimensionless quantities, e.g. mole fraction or mass fraction. Since these fractions are quantity-per-quantity measures, they ...
of blood in seawater. Sharks have the ability to determine the direction of a given scent based on the timing of scent detection in each nostril. This is similar to the method mammals use to determine direction of sound. They are more attracted to the chemicals found in the intestines of many species, and as a result often linger near or in
sewage Sewage (or domestic sewage, domestic wastewater, municipal wastewater) is a type of wastewater that is produced by a community of people. It is typically transported through a sewer system. Sewage consists of wastewater discharged from residenc ...
outfalls. Some species, such as
nurse shark The nurse shark (''Ginglymostoma cirratum'') is an elasmobranch fish in the family Ginglymostomatidae. The conservation status of the nurse shark is globally assessed as Vulnerable in the IUCN List of Threatened Species. They are considered t ...
s, have external
barbels In fish anatomy and turtle anatomy, a barbel is a slender, whiskerlike sensory organ near the mouth. Fish that have barbels include the catfish, the carp, the goatfish, the hagfish, the sturgeon, the zebrafish, the black dragonfish and some ...
that greatly increase their ability to sense prey.


Magnetoception


Electroreception

Some fish, such as catfish and sharks, have organs that detect weak electric currents on the order of millivolt. Other fish, like the South American electric fishes
Gymnotiformes The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water (the ...
, can produce weak electric currents, which they use in navigation and social communication. In sharks, the
ampullae of Lorenzini Ampullae of Lorenzini (singular ''Ampulla'') are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of cartilaginous fish (sharks, rays, and chimaeras) and of basal actinopterygia ...
are electroreceptor organs. They number in the hundreds to thousands. Sharks use the ampullae of Lorenzini to detect the
electromagnetic field An electromagnetic field (also EM field or EMF) is a classical (i.e. non-quantum) field produced by (stationary or moving) electric charges. It is the field described by classical electrodynamics (a classical field theory) and is the classical c ...
s that all living things produce. This helps sharks (particularly the
hammerhead shark The hammerhead sharks are a group of sharks that form the family Sphyrnidae, so named for the unusual and distinctive structure of their heads, which are flattened and laterally extended into a "hammer" shape called a cephalofoil. Most hammerhe ...
) find prey. The shark has the greatest electrical sensitivity of any animal. Sharks find prey hidden in sand by detecting the
electric field An electric field (sometimes E-field) is the physical field that surrounds electrically charged particles and exerts force on all other charged particles in the field, either attracting or repelling them. It also refers to the physical field fo ...
s they produce.
Ocean current An ocean current is a continuous, directed movement of sea water generated by a number of forces acting upon the water, including wind, the Coriolis effect, breaking waves, cabbeling, and temperature and salinity differences. Depth contours, s ...
s moving in the
magnetic field of the Earth Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from the Sun. The magnetic fi ...
also generate electric fields that sharks can use for orientation and possibly navigation. * The
ampullae of Lorenzini Ampullae of Lorenzini (singular ''Ampulla'') are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of cartilaginous fish (sharks, rays, and chimaeras) and of basal actinopterygia ...
allow sharks to sense electrical discharges. *
Electric fish An electric fish is any fish that can generate electric fields. Most electric fish are also electroreceptive, meaning that they can sense electric fields. The only exception is the stargazer family. Electric fish, although a small minority, in ...
are able to produce electric fields by modified muscles in their body.


Pain

Experiments done by William Tavolga provide evidence that fish have
pain Pain is a distressing feeling often caused by intense or damaging stimuli. The International Association for the Study of Pain defines pain as "an unpleasant sensory and emotional experience associated with, or resembling that associated with, ...
and fear responses. For instance, in Tavolga's experiments,
toadfish Toadfish is the common name for a variety of species from several different families of fish, usually because of their toad-like appearance. "Dogfish" is a name for certain species along the gulf coast. Dolphin-Toadfish relationship Toadfish mak ...
grunted when electrically shocked and over time they came to grunt at the mere sight of an electrode. In 2003, Scottish scientists at the
University of Edinburgh The University of Edinburgh ( sco, University o Edinburgh, gd, Oilthigh Dhùn Èideann; abbreviated as ''Edin.'' in post-nominals) is a public research university based in Edinburgh, Scotland. Granted a royal charter by King James VI in 15 ...
and the Roslin Institute concluded that rainbow trout exhibit behaviors often associated with
pain Pain is a distressing feeling often caused by intense or damaging stimuli. The International Association for the Study of Pain defines pain as "an unpleasant sensory and emotional experience associated with, or resembling that associated with, ...
in other animals. Bee
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
and
acetic acid Acetic acid , systematically named ethanoic acid , is an acidic, colourless liquid and organic compound with the chemical formula (also written as , , or ). Vinegar is at least 4% acetic acid by volume, making acetic acid the main component ...
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. Neurons fired in a pattern resembling human neuronal patterns. Professor James D. Rose of the
University of Wyoming The University of Wyoming (UW) is a public land-grant research university in Laramie, Wyoming. It was founded in March 1886, four years before the territory was admitted as the 44th state, and opened in September 1887. 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". 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 The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
. However, animal behaviorist
Temple Grandin Mary Temple Grandin (born August 29, 1947) is an American academic and animal behaviorist. She is a prominent proponent for the humane treatment of livestock for slaughter and the author of more than 60 scientific papers on animal behavior. Gra ...
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." Animal welfare advocates raise concerns about the possible
suffering Suffering, or pain in a broad sense, may be an experience of unpleasantness or aversion, possibly associated with the perception of harm or threat of harm in an individual. Suffering is the basic element that makes up the negative valence of a ...
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.


Reproductive processes

Oogonia An oogonium (plural oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in larg ...
development in teleosts fish varies according to the group, and the determination of oogenesis dynamics allows the understanding of maturation and fertilisation processes. Changes in the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...
, ooplasm, and the surrounding layers characterize the oocyte maturation process. Postovulatory follicles are structures formed after oocyte release; they do not have
endocrine The endocrine system is a messenger system comprising feedback loops of the hormones released by internal glands of an organism directly into the circulatory system, regulating distant target organs. In vertebrates, the hypothalamus is the neu ...
function, present a wide irregular lumen, and are rapidly reabsorbed in a process involving the
apoptosis Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes (morphology) and death. These changes incl ...
of follicular cells. A degenerative process called
follicular atresia Follicular atresia refers to the process in which a follicle fails to develop, thus preventing it from ovulating and releasing an egg. It is a normal, naturally occurring progression that occurs as mammalian ovaries age. Approximately 1% of mamma ...
reabsorbs vitellogenic oocytes not spawned. This process can also occur, but less frequently, in oocytes in other development stages. Some fish are
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
s, having both testes and ovaries either at different phases in their life cycle or, as in
hamlets A hamlet is a human settlement that is smaller than a town or village. Its size relative to a parish can depend on the administration and region. A hamlet may be considered to be a smaller settlement or subdivision or satellite entity to a lar ...
, have them simultaneously. Over 97% of all known fish are
oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and ...
,Peter Scott: ''Livebearing Fishes'', p. 13. Tetra Press 1997. that is, the eggs develop outside the mother's body. Examples of oviparous fish include
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
,
goldfish The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the wild have ...
,
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
s,
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
, and
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s. In the majority of these species, fertilisation takes place outside the mother's body, with the male and female fish shedding their
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s into the surrounding water. However, a few oviparous fish practice internal fertilisation, with the male using some sort of
intromittent organ An intromittent organ is any external organ of a male organism that is specialized to deliver sperm during copulation. Intromittent organs are found most often in terrestrial species, as most non-mammalian aquatic species fertilize their eggs e ...
to deliver sperm into the genital opening of the female, most notably the oviparous sharks, such as the
horn shark The horn shark (''Heterodontus francisci'') is a species of bullhead shark, in the family Heterodontidae. It is endemic to the coastal waters off the western coast of North America, from California to the Gulf of California. Young sharks are seg ...
, and oviparous rays, such as skates. In these cases, the male is equipped with a pair of modified
pelvic The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
fins known as
claspers In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's ...
. Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of . The eggs are generally surrounded by the extraembryonic membranes but do not develop a shell, hard or soft, around these membranes. Some fish have thick, leathery coats, especially if they must withstand physical force or desiccation. These type of eggs can also be very small and fragile. File:Oeufs002b,57.png, Egg of
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
File:Oeufs002b,54.png, Egg of
catshark Catsharks are ground sharks of the family Scyliorhinidae. They are the largest family of sharks with around 160 species placed in 17 genera. Although they are generally known as catsharks, some species can also be called dogfish due to previous n ...
( mermaids' purse) File:Oeufs002b,55.png, Egg of bullhead shark File:Oeufs002b,56.png, Egg of
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
The newly hatched young of oviparous fish are called
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e. They are usually poorly formed, carry a large
yolk sac The yolk sac is a membranous sac attached to an embryo, formed by cells of the hypoblast layer of the bilaminar embryonic disc. This is alternatively called the umbilical vesicle by the Terminologia Embryologica (TE), though ''yolk sac'' is far ...
(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 Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
prey, a process which depends on typically inadequate zooplankton density, starving many larvae. In
ovoviviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
fish the eggs develop inside the mother's body after internal fertilisation but receive little or no nourishment directly from the mother, depending instead on the
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example bec ...
. Each embryo develops in its own egg. Familiar examples of ovoviviparous fish include
guppies The guppy (), also known as millionfish and rainbow fish, is one of the world's most widely distributed tropical fish and one of the most popular List of freshwater aquarium fish species, freshwater aquarium fish species. It is a member of the ...
,
angel shark The angelsharks are a group of sharks in the genus ''Squatina'' of the family Squatinidae. They commonly inhabit sandy seabeds close to in depth. Many species are now classified as critically endangered by the International Union for Conservat ...
s, and
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
s. Some species of fish are
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
. In such species the mother retains the eggs and nourishes the embryos. Typically, viviparous fish have a structure analogous to the
placenta The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mater ...
seen in
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
connecting the mother's blood supply with that of the embryo. Examples of viviparous fish include the surf-perches, splitfins, and
lemon shark The lemon shark (''Negaprion brevirostris'') is a species of shark from the family Carcharhinidae and is classified as a Vulnerable species by the International Union for the Conservation of Nature. Lemon sharks can grow to in length. They are o ...
. Some viviparous fish exhibit
oophagy Oophagy ( ) sometimes ovophagy, literally "egg eating", is the practice of embryos feeding on eggs produced by the ovary while still inside the mother's uterus. The word oophagy is formed from the classical Greek (, "egg") and classical Greek (, ...
, in which the developing embryos eat other eggs produced by the mother. This has been observed primarily among sharks, such as the
shortfin mako The shortfin mako shark (; ; ''Isurus oxyrinchus''), also known as the blue pointer or bonito shark, is a large Lamniformes, mackerel shark. It is commonly referred to as the mako shark, as is the longfin mako shark (''Isurus paucus''). The shor ...
and
porbeagle The porbeagle (''Lamna nasus'') is a species of mackerel shark in the family Lamnidae, distributed widely in the cold and temperate marine waters of the North Atlantic and Southern Hemisphere. In the North Pacific, its ecological equivalent is ...
, but is known for a few bony fish as well, such as the
halfbeak Hemiramphidae is a family of fishes that are commonly called halfbeaks, spipe fish or spipefish. They are a geographically widespread and numerically abundant family of epipelagic fish inhabiting warm waters around the world. The halfbeaks are ...
''Nomorhamphus ebrardtii''.
Intrauterine cannibalism Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is well documented, bo ...
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 The sand tiger shark (''Carcharias taurus''), gray nurse shark, spotted ragged-tooth shark or blue-nurse sand tiger, is a species of shark that inhabits subtropical and temperate waters worldwide. It inhabits the continental shelf, from sandy sho ...
, but has also been reported for ''Nomorhamphus ebrardtii''. In many species of fish, fins have been modified to allow
Internal fertilisation Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For inter ...
.
Aquarists Fishkeeping is a popular hobby, practiced by aquarists, concerned with keeping fish in a home aquarium or garden pond. There is also a piscicultural fishkeeping industry, serving as a branch of agriculture. Origins of fishkeeping Fish have ...
commonly refer to ovoviviparous and viviparous fish as
livebearers Livebearers are aquarium fish that retain the eggs inside the body and give birth to live, free-swimming young. Among aquarium fish, livebearers are nearly all members of the family Poeciliidae and include guppies, mollies, platies and swordta ...
. * Many fish species are
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
s. ''Synchronous hermaphrodites'' possess both
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
and
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoster ...
at the same time. ''Sequential hermaphrodites'' have both types of tissue in their
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s, with one type being predominant while the fish belongs to the corresponding gender.


Social behaviour

Fish social behaviour called ‘shoaling’ involves a group of fish swimming together. This behaviour is a defence mechanism in the sense that there is safety in large numbers, where chances of being eaten by predators are reduced. Shoaling also increases mating and foraging success. Schooling on the other hand, is a behaviour within the shoal where fish can be seen performing various manoeuvres in a synchronised manner. The parallel swimming is a form of ‘social copying’ where fish in the school replicate the direction and velocity of its neighbouring fishes. Experiments done by D.M. Steven, on the shoaling behaviour of fish concluded that during the day, fish had a higher tendency to stay together as a result of a balance between single fish leaving and finding their own direction and the mutual attraction between fishes of the same species. It was found that at night the fish swam noticeably faster however, often singly and in no co-ordination. Groups of two or three could be seen frequently formed although were dispersed after a couple of seconds. Theoretically, the amount of time that a fish stays together in a shoal should represent their cost of staying instead of leaving. A past laboratory experiment done on cyprinids has established that the time budget for social behaviour within a shoal varies proportionally to the quantity of fishes present. This originates from the cost/benefit ratio which changes accordingly with group size, measured by the risk of predation versus food intake. When the cost/benefit ratio is favourable to shoaling behaviour then decisions to stay with a group or join one is favourable. Depending on this ratio, fish will correspondingly decide to leave or stay. Thus, shoaling behaviour is considered to be driven by an individual fish's constant stream of decisions.


See also

*
Anatomical terms of location Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
* Digital Fish Library *
Evolution of fish 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 anatomy 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 co ...
* Fish development *
Fish measurement Fish measurement is the measuring of individual fish and various parts of their anatomies. These data are used in many areas of ichthyology, including taxonomy and fisheries biology. Overall length * Standard length (SL) is the length of a fish ...
*
Ichthyology terms This glossary of ichthyology is a list of definitions of terms and concepts used in ichthyology, the study of fishes. A B C ...
* Panderichthys digits


References


Further reading

* Bernier NJ, Van Der Kraak G, Farrell AP and Brauner CJ (2009
''Fish Physiology: Fish Neuroendocrinology''
Academic Press. . * Eddy FB and Handy RD (2012
'' Ecological and Environmental Physiology of Fishes''
Oxford University Press. . * Evans DH, JB Claiborne and S Currie (Eds) (2013
''The Physiology of Fishes''
4th edition, CRC Press. . * Grosell M, Farrell AP and Brauner CJ (2010
''Fish Physiology: The Multifunctional Gut of Fish''
Academic Press. . * Hara TJ and Zielinski B (2006
''Fish Physiology: Sensory Systems Neuroscience''
Academic Press. . * Kapoor BG and Khanna B (2004
"Ichthyology handbook"
Pages 137–140, Springer. . * McKenzie DJ, Farrell AP and Brauner CJ (2007
''Fish Physiology: Primitive Fishes''
Academic Press. . * Sloman KA, Wilson RW and Balshine S (2006
''Behaviour And Physiology of Fish''
Gulf Professional Publishing. . * Wood CM, Farrell AP and Brauner CJ (2011
''Fish Physiology: Homeostasis and Toxicology of Non-Essential Metals''
Academic Press. .


External links

{{diversity of fish