Fish physiology
Fish physiology is the scientific study of how the component parts of fish function together in the living fish.[2] It can be contrasted with fish anatomy, which is the study of the form or morphology of fishes. In practice, fish anatomy and physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the later dealing with how those components function together in the living fish. For this, at first we need to know about their intestinal morphology.
Respiration
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Most fish exchange gases using gills on either side of the pharynx (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.[3][4] Each filament contains a capillary network that provides a large surface area for exchanging oxygen and carbon dioxide. Fish exchange gases by pulling oxygen-rich water through their mouths and pumping it over their gills. In some fish, capillary blood flows in the opposite direction to the water, causing countercurrent exchange. 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.
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 cichlids, quickly suffocate, while air-breathers survive for much longer, in some cases in water that is little more than wet mud. At the most extreme, some air-breathing fish are able to survive in damp burrows for weeks without water, entering a state of aestivation (summertime hibernation) until water returns.
Air breathing fish can be divided into obligate air breathers and facultative air breathers. Obligate air breathers, such as the
All
The gills are composed of comb-like filaments, the
Scientists have investigated what part of the body is responsible for maintaining the respiratory rhythm. They found that
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. The mechanisms by which these changes occur have been strongly debated over more than 100 years between scientists.[13] 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 centers of the brain connect to respiratory centers 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, 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.[11]
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.[11]
Marine
In some primitive bony fishes and
Cartilaginous fish
Like other fish, sharks extract
The respiration and circulation process begins when deoxygenated blood travels to the shark's two-chambered heart. Here the shark pumps blood to its gills via the ventral aorta artery 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 blood flows from the dorsal aorta throughout the body. The deoxygenated blood from the body then flows through the posterior cardinal veins and enters the posterior cardinal sinuses. From there blood enters the heart ventricle and the cycle repeats.[18]
A smaller opening, the spiracle, lies in the back of the first gill slit. This bears a small pseudobranch that resembles a gill in structure, but only receives blood already oxygenated by the true gills.[11] The spiracle is thought to be homologous to the ear opening in higher vertebrates.[19]
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.[11]
Chimaeras differ from other cartilagenous fish, having lost both the spiracle and the fifth gill slit. The remaining slits are covered by an operculum, developed from the septum of the gill arch in front of the first gill.[11]
Lampreys and hagfish
Lampreys and hagfish do not have gill slits as such. Instead, the gills are contained in spherical pouches, with a circular opening to the outside. Like the gill slits of higher fish, each pouch contains two gills. In some cases, the openings may be fused together, effectively forming an operculum. Lampreys have seven pairs of pouches, while hagfishes may have six to fourteen, depending on the species. In the hagfish, the pouches connect with the pharynx internally. In adult lampreys, a separate respiratory tube develops beneath the pharynx proper, separating food and water from respiration by closing a valve at its anterior end.[11]
Circulation
The circulatory systems of all
In amphibians and most reptiles, a
Digestion
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
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, lampreys, hagfishes, chimaeras, lungfishes, and some teleost 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.[11]
The small intestine is the part of the digestive tract following the stomach and followed by the large intestine, and is where much of the digestion 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.[21] The small intestine is found in all teleosts, 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 being marked only by the end of the digestive epithelium.[11]
There is no small intestine as such in non-teleost fish, such as sharks, sturgeons, and lungfish. 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.[11] In lampreys, the spiral valve is extremely small, possibly because their diet requires little digestion. Hagfish have no spiral valve at all, with digestion occurring for almost the entire length of the intestine, which is not subdivided into different regions.[11]
The
As with many aquatic animals, most fish release their nitrogenous wastes as
Saltwater fish tend to lose water because of osmosis. Their kidneys return water to the body. The reverse happens in freshwater fish: they tend to gain water osmotically. Their kidneys produce dilute urine for excretion. Some fish have specially adapted kidneys that vary in function, allowing them to move from freshwater to saltwater.
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.[23] 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 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.[23]
Endocrine system
Regulation of social behaviour
Oxytocin is a group of neuropeptides found in most vertebrates. One form of oxytocin functions as a hormone which is associated with human love. In 2012, researchers injected cichlids from the social species Neolamprologus pulcher, 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".[24][25]
Effects of pollution
Fish can
Freshwater habitats in the United States are widely contaminated by the common pesticide atrazine.[29] There is controversy over the degree to which this pesticide harms the endocrine systems of freshwater fish and amphibians. Non-industry-funded researchers consistently report harmful effects while industry-funded researchers consistently report no harmful effects.[29][30][31]
In the marine ecosystem,
- 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 are osmoconformers and osmoregulators. Osmoconformers 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.
In contrast to bony fish, with the exception of the
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
Thermoregulation
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 at which an organism can survive indefinitely.[38]
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
To cope with low temperatures, some
Most sharks are "cold-blooded" or, more precisely,
- porbeagle sharks who maintain body temperatures elevated in excess of 20 °C (36 °F) above ambient water temperatures.[42] See also gigantothermy. Endothermy, though metabolically costly, is thought to provide advantages such as increased muscle strength, higher rates of central nervous system processing, and higher rates of digestion.
In some fish, a
The eye of a
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. 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.[44]
Mostly fish have white muscles, but the muscles of some fishes, such as
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
A typical characteristic of many animals that utilize undulatory locomotion is that they have segmented muscles, or blocks of myomeres, 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.[45] 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.[46] When a pennate muscle 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).[46] 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).[46]
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
Buoyancy
The body of a fish is denser than water, so fish must compensate for the difference or they will sink. Many
In some fish, a rete mirabile fills the swim bladder with oxygen. A
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
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
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.[54]
Vision
Hearing
Fish can sense sound through their lateral lines and their otoliths (ears). Some fishes, such as some species of carp and herring, hear through their swim bladders, which function rather like a hearing aid.[58]
Hearing is well-developed in
Although it is hard to test sharks' hearing, they may have a sharp
Chemoreception
Sharks have keen
Sharks have the ability to determine the direction of a given scent based on the timing of scent detection in each nostril.[61] 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
Magnetoception
Electroreception
Some fish, such as catfish and sharks, have organs that detect weak electric currents on the order of millivolt.[62] Other fish, like the South American electric fishes Gymnotiformes, can produce weak electric currents, which they use in navigation and social communication. In sharks, the ampullae of Lorenzini are electroreceptor organs. They number in the hundreds to thousands. Sharks use the ampullae of Lorenzini to detect the electromagnetic fields that all living things produce.[63] This helps sharks (particularly the hammerhead shark) find prey. The shark has the greatest electrical sensitivity of any animal. Sharks find prey hidden in sand by detecting the electric fields they produce. Ocean currents moving in the magnetic field of the Earth also generate electric fields that sharks can use for orientation and possibly navigation.[64]
- The ampullae of Lorenzini allow sharks to sense electrical discharges.
- Electric fish are able to produce electric fields by modified muscles in their body.
Pain
Experiments done by William Tavolga provide evidence that fish have pain and fear responses. For instance, in Tavolga's experiments, toadfish grunted when electrically shocked and over time they came to grunt at the mere sight of an electrode.[65]
In 2003, Scottish scientists at the University of Edinburgh and the Roslin Institute concluded that rainbow trout exhibit behaviors often associated with pain in other animals. Bee venom and acetic acid injected into the lips resulted in fish rocking their bodies and rubbing their lips along the sides and floors of their tanks, which the researchers concluded were attempts to relieve pain, similar to what mammals would do.[66][67][68] Neurons fired in a pattern resembling human neuronal patterns.[68]
Professor James D. Rose of the University of Wyoming claimed the study was flawed since it did not provide proof that fish possess "conscious awareness, particularly a kind of awareness that is meaningfully like ours".[69] 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.[70] However, animal behaviorist Temple Grandin argues that fish could still have consciousness without a neocortex because "different species can use different brain structures and systems to handle the same functions."[68]
Animal welfare advocates raise concerns about the possible suffering of fish caused by angling. Some countries, such as Germany have banned specific types of fishing, and the British RSPCA now formally prosecutes individuals who are cruel to fish.[71]
Reproductive processes
Postovulatory
Some fish are
Over 97% of all known fish are
Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of 1 millimetre (0.039 in). 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.
-
Egg of lamprey
-
Egg ofmermaids' purse)
-
Egg of bullhead shark
-
Egg of chimaera
The newly hatched young of oviparous fish are called larvae. They are usually poorly formed, carry a large yolk sac (for nourishment) and are very different in appearance from juvenile and adult specimens. The larval period in oviparous fish is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on zooplankton prey, a process which depends on typically inadequate zooplankton density, starving many larvae.
In
Some species of fish are
In many species of fish, fins have been modified to allow Internal fertilisation.
Aquarists commonly refer to ovoviviparous and viviparous fish as livebearers.
- Many fish species are hermaphrodites. Synchronous hermaphrodites possess both ovaries and testes at the same time. Sequential hermaphrodites have both types of tissue in their gonads, 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.[75] The parallel swimming is a form of ‘social copying’ where fish in the school replicate the direction and velocity of its neighbouring fishes.[76]
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.[77]
Theoretically, the amount of time that a fish stays together in a shoal should represent their cost of staying instead of leaving.[78] 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.[79] 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.[75]
See also
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Further reading
- Bernier NJ, Van Der Kraak G, Farrell AP and Brauner CJ (2009) Fish Physiology: Fish Neuroendocrinology Academic Press. ISBN 978-0-08-087798-3.
- Eddy FB and Handy RD (2012) Ecological and Environmental Physiology of Fishes Oxford University Press. ISBN 978-0-19-954095-2.
- Evans DH, JB Claiborne and S Currie (Eds) (2013) The Physiology of Fishes 4th edition, CRC Press. ISBN 978-1-4398-8030-2.
- Grosell M, Farrell AP and Brauner CJ (2010) Fish Physiology: The Multifunctional Gut of Fish Academic Press. ISBN 978-0-08-096136-1.
- Hara TJ and Zielinski B (2006) Fish Physiology: Sensory Systems Neuroscience Academic Press. ISBN 978-0-08-046961-4.
- Kapoor BG and Khanna B (2004) "Ichthyology handbook" Pages 137–140, Springer. ISBN 978-3-540-42854-1.
- McKenzie DJ, Farrell AP and Brauner CJ (2007) Fish Physiology: Primitive Fishes Academic Press. ISBN 978-0-08-054952-1.
- Sloman KA, Wilson RW and Balshine S (2006) Behaviour And Physiology of Fish Gulf Professional Publishing. ISBN 978-0-12-350448-7.
- Wood CM, Farrell AP and Brauner CJ (2011) Fish Physiology: Homeostasis and Toxicology of Non-Essential Metals Academic Press. ISBN 978-0-12-378634-0.