Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

“All religions, arts and sciences are branches of the same tree. All these aspirations are directed toward ennobling man's life, lifting it from the sphere of mere physical existence and leading the individual towards freedom.”—Albert Einstein

 

“Mathematics, rightly viewed, possesses not only truth, but supreme beauty—a beauty cold and austere, like that of sculpture, without appeal to any part of our weaker nature, without the gorgeous trappings of painting or music, yet sublimely pure, and capable of a stern perfection such as only the greatest art can show.” –Bertrand Russell, “A History of Western Philosophy”

 

“Sculpture is the art of intelligence.”—Pablo Picasso

 

“[T]here shall be love between the poet and the man of demonstrable science. In the beauty of poems are the tuft and final applause of science.”—Walt Whitman

Dwight D. Eisenhower

My fellow Americans:

Three days from now, after half a century in the service of our country, I shall lay down the responsibilities of office as, in traditional and solemn ceremony, the authority of the Presidency is vested in my successor.

This evening I come to you with a message of leave-taking and farewell, and to share a few final thoughts with you, my countrymen.

Like every other citizen, I wish the new President, and all who will labor with him, Godspeed. I pray that the coming years will be blessed with peace and prosperity for all.

Our people expect their President and the Congress to find essential agreement on issues of great moment, the wise resolution of which will better shape the future of the Nation.

My own relations with the Congress, which began on a remote and tenuous basis when, long ago, a member of the Senate appointed me to West Point, have since ranged to the intimate during the war and immediate post-war period, and, finally, to the mutually interdependent during these past eight years.

In this final relationship, the Congress and the Administration have, on most vital issues, cooperated well, to serve the national good rather than mere partisanship, and so have assured that the business of the Nation should go forward. So, my official relationship with the Congress ends in a feeling, on my part, of gratitude that we have been able to do so much together.

II.

We now stand ten years past the midpoint of a century that has witnessed four major wars among great nations. Three of these involved our own country. Despite these holocausts America is today the strongest, the most influential and most productive nation in the world. Understandably proud of this pre-eminence, we yet realize that America's leadership and prestige depend, not merely upon our unmatched material progress, riches and military strength, but on how we use our power in the interests of world peace and human betterment.

III.

Throughout America's adventure in free government, our basic purposes have been to keep the peace; to foster progress in human achievement, and to enhance liberty, dignity and integrity among people and among nations. To strive for less would be unworthy of a free and religious people. Any failure traceable to arrogance, or our lack of comprehension or readiness to sacrifice would inflict upon us grievous hurt both at home and abroad.

Progress toward these noble goals is persistently threatened by the conflict now engulfing the world. It commands our whole attention, absorbs our very beings. We face a hostile ideology -- global in scope, atheistic in character, ruthless in purpose, and insidious in method. Unhappily the danger is poses promises to be of indefinite duration. To meet it successfully, there is called for, not so much the emotional and transitory sacrifices of crisis, but rather those which enable us to carry forward steadily, surely, and without complaint the burdens of a prolonged and complex struggle -- with liberty the stake. Only thus shall we remain, despite every provocation, on our charted course toward permanent peace and human betterment.

Crises there will continue to be. In meeting them, whether foreign or domestic, great or small, there is a recurring temptation to feel that some spectacular and costly action could become the miraculous solution to all current difficulties. A huge increase in newer elements of our defense; development of unrealistic programs to cure every ill in agriculture; a dramatic expansion in basic and applied research -- these and many other possibilities, each possibly promising in itself, may be suggested as the only way to the road we wish to travel.

But each proposal must be weighed in the light of a broader consideration: the need to maintain balance in and among national programs -- balance between the private and the public economy, balance between cost and hoped for advantage -- balance between the clearly necessary and the comfortably desirable; balance between our essential requirements as a nation and the duties imposed by the nation upon the individual; balance between actions of the moment and the national welfare of the future. Good judgment seeks balance and progress; lack of it eventually finds imbalance and frustration.

The record of many decades stands as proof that our people and their government have, in the main, understood these truths and have responded to them well, in the face of stress and threat. But threats, new in kind or degree, constantly arise. I mention two only.

IV.

A vital element in keeping the peace is our military establishment. Our arms must be mighty, ready for instant action, so that no potential aggressor may be tempted to risk his own destruction.

Our military organization today bears little relation to that known by any of my predecessors in peacetime, or indeed by the fighting men of World War II or Korea.

Until the latest of our world conflicts, the United States had no armaments industry. American makers of plowshares could, with time and as required, make swords as well. But now we can no longer risk emergency improvisation of national defense; we have been compelled to create a permanent armaments industry of vast proportions. Added to this, three and a half million men and women are directly engaged in the defense establishment. We annually spend on military security more than the net income of all United States corporations.

This conjunction of an immense military establishment and a large arms industry is new in the American experience. The total influence -- economic, political, even spiritual -- is felt in every city, every State house, every office of the Federal government. We recognize the imperative need for this development. Yet we must not fail to comprehend its grave implications. Our toil, resources and livelihood are all involved; so is the very structure of our society.

In the councils of government, we must guard against the acquisition of unwarranted influence, whether sought or unsought, by the militaryindustrial complex. The potential for the disastrous rise of misplaced power exists and will persist.

We must never let the weight of this combination endanger our liberties or democratic processes. We should take nothing for granted. Only an alert and knowledgeable citizenry can compel the proper meshing of the huge industrial and military machinery of defense with our peaceful methods and goals, so that security and liberty may prosper together.

Akin to, and largely responsible for the sweeping changes in our industrial-military posture, has been the technological revolution during recent decades.

In this revolution, research has become central; it also becomes more formalized, complex, and costly. A steadily increasing share is conducted for, by, or at the direction of, the Federal government.

Today, the solitary inventor, tinkering in his shop, has been overshadowed by task forces of scientists in laboratories and testing fields. In the same fashion, the free university, historically the fountainhead of free ideas and scientific discovery, has experienced a revolution in the conduct of research. Partly because of the huge costs involved, a government contract becomes virtually a substitute for intellectual curiosity. For every old blackboard there are now hundreds of new electronic computers.

The prospect of domination of the nation's scholars by Federal employment, project allocations, and the power of money is ever present

•and is gravely to be regarded.

Yet, in holding scientific research and discovery in respect, as we should, we must also be alert to the equal and opposite danger that public policy could itself become the captive of a scientifictechnological elite.

It is the task of statesmanship to mold, to balance, and to integrate these and other forces, new and old, within the principles of our democratic system -- ever aiming toward the supreme goals of our free society.

V.

Another factor in maintaining balance involves the element of time. As we peer into society's future, we -- you and I, and our government -- must avoid the impulse to live only for today, plundering, for our own ease and convenience, the precious resources of tomorrow. We cannot mortgage the material assets of our grandchildren without risking the loss also of their political and spiritual heritage. We want democracy to survive for all generations to come, not to become the insolvent phantom of tomorrow.

VI.

Down the long lane of the history yet to be written America knows that this world of ours, ever growing smaller, must avoid becoming a community of dreadful fear and hate, and be instead, a proud confederation of mutual trust and respect.

Such a confederation must be one of equals. The weakest must come to the conference table with the same confidence as do we, protected as we are by our moral, economic, and military strength. That table, though scarred by many past frustrations, cannot be abandoned for the certain agony of the battlefield.

Disarmament, with mutual honor and confidence, is a continuing imperative. Together we must learn how to compose differences, not with arms, but with intellect and decent purpose. Because this need is so sharp and apparent I confess that I lay down my official responsibilities in this field with a definite sense of disappointment. As one who has witnessed the horror and the lingering sadness of war -- as one who knows that another war could utterly destroy this civilization which has been so slowly and painfully built over thousands of years -- I wish I could say tonight that a lasting peace is in sight.

Happily, I can say that war has been avoided. Steady progress toward our ultimate goal has been made. But, so much remains to be done. As a private citizen, I shall never cease to do what little I can to help the world advance along that road.

VII.

So -- in this my last good night to you as your President -- I thank you for the many opportunities you have given me for public service in war and peace. I trust that in that service you find some things worthy; as for the rest of it, I know you will find ways to improve performance in the future.

You and I -- my fellow citizens -- need to be strong in our faith that all nations, under God, will reach the goal of peace with justice. May we be ever unswerving in devotion to principle, confident but humble with power, diligent in pursuit of the Nation's great goals.

To all the peoples of the world, I once more give expression to America's prayerful and continuing aspiration:

We pray that peoples of all faiths, all races, all nations, may have their great human needs satisfied; that those now denied opportunity shall come to enjoy it to the full; that all who yearn for freedom may experience its spiritual blessings; that those who have freedom will understand, also, its heavy responsibilities; that all who are insensitive to the needs of others will learn charity; that the scourges of poverty, disease and ignorance will be made to disappear from the earth, and that, in the goodness of time, all peoples will come to live together in a peace guaranteed by the binding force of mutual respect and love.

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Abstract: Sustainability has the potential to provide a holistic framework that can bridge the gap that is often found between socio-economic justice and environmental discourses. However, sustainability and sustainability education have typically accepted the prevailing socio-economic and cultural paradigm. It is my aim in this paper to demonstrate that a truly holistic and visionary sustainability (education) framework ought to demand radical and critical theories and solutions- based approaches to politicize and interrogate the premises, assumptions, and biases linked to the dominant notion of sustainability. If we are to envision and construe actual sustainable futures, we must first understand what brought us here, where the roots of the problems lie, and how the sustainability discourse and framework tackle—or fail to tackle—them. To do this is to politicize sustainability, to build a critical perspective of and about sustainability. It is an act of conscientização (or conscientization), to borrow Paulo Freire’s seminal term, of cultivating critical consciousness and conscience. In lieu of the standard articulation of politics as centralized state administration, ‘critical sustainability studies’ is based on a framing that gives prominence to a more organic, decentralized engagement of conscientious subjects in the creation of just, regenerative eco-social relations. It illuminates the ideological and material links between society, culture, and ecology by devoting particular attention to how knowledge and discourse around and across those realms are generated and articulated. I believe that future scholarship and activism in sustainability and sustainability-related fields would benefit immensely from dialoguing with this framework.

 

The assumption that what currently exists must necessarily exist is the acid that corrodes all visionary thinking.

 

– Murray Bookchin, The Meaning of Confederalism, 1990

   

Introduction: Why Sustainability (and Sustainability Education)?

 

Despite conflicting opinions over what the terms ‘sustainability’ and its variant ‘sustainable development’ actually mean, the framework of sustainability has gained a lot of traction in the last two decades. Its Western origins can be traced back to the writings of Western philosophers and seminal environmentalists like John Locke and Aldo Leopold (Spoon, 2013). Redclift (2005) asserts that sustainability as an idea was first used during the ‘limits to growth’ debates in the 1970s and the 1972 UN Stockholm Conference. Perhaps the most commonly quoted definition of sustainable development is that of the World Commission on Environment and Development (WCED) who states that “sustainable development is development that meets the needs of the present without compromising the ability of future generations to meet their own needs” (WCED, 1987, p. 43).

 

Sustainability has the potential to provide a holistic framework that can bridge the gap that is often found between socio-economic justice and environmental discourses. After all, recent scholarship indicates that the issue of environmental quality is inevitably linked to that of human equity (Morello-Frosch, 1997; Torras & Boyce, 1998; see Agyeman, Bullard, & Evans, 2002), and thus they need to be thought about together. I hold that an actual sustainable society is one where wider matters of social and economic needs are intrinsically connected to the dynamic limits set by supporting ecosystems and environments.

 

Sustainability education has emerged as an effort to acknowledge and reinforce these interrelationships and to reorient and transform education along the lines of social and ecological well-being (Sterling, 2001). By being rooted in whole systems thinking, i.e. “the ability to collectively analyze complex systems across different domains (society, environment, and economy) and across different scales (local to global)” (Wiek, Withycombe, & Redman, 2011, p. 207), sustainability education strives to illuminate the complexities associated with the broad, problem-oriented, solution-driven nature of sustainability (Warren, Archambault, & Foley, 2014). If we are to devise cultural systems that are truly regenerative, this “novel” brand of education urges the teaching of the fundamental facts of life by stewarding learning communities that comprehend the adaptive qualities of ecological patterns and principles (Stone, 2012). Sustainability education highlights the centrality of ‘place’ as a unit of inquiry to devise reciprocal—and thus sustainable—relationships where one nourishes and is nourished by their surrounding social and ecological milieus (Williams & Brown, 2012).

 

Additionally, sustainability and, as a consequence, sustainability education are future- oriented and therefore demand ‘futures thinking’: the ability to assess and formulate nuanced pictures of the future vis-à-vis sustainability predicaments and sustainability problem-solving schemes (Wiek, et al., 2011). In a nutshell, futures thinking suggests that we need to imagine the potential ramifications of past and current human activities by critically analyzing them today if we are to conceive of new, more sustainable futures (Warren et al., 2014). Future studies can therefore help people to pursue their “ontological vocation” as history makers (Freire, 1993, p. 66) and to (re)claim their agency as a means of creating the world in which they wish to live (Inayatullah, 2007).

 

However, sustainability and sustainability education have typically accepted the prevailing socio-economic and cultural paradigm despite their apparent holistic intent and(theoretical) efforts to reconcile the three pillars of sustainability—equity, environment, and economy. Whether intentionally or not, they have promoted curative solutions instead of reflecting new, critical mindsets that can actually generate meaningful socio-cultural innovation by naming and discursively dismantling the systems and processes that are the root causes of the complex problems we face. And, as Albert Einstein once put it, “no problem can be solved from the same consciousness that created it.”

 

It is my aim in this paper to demonstrate that a truly holistic and visionary sustainability (education) framework ought to demand radical (of, relating to, or proceeding from a root) and critical (of, relating to, or being a turning point) theories and solutions-based approaches to politicize and interrogate the premises, assumptions, and biases linked to the dominant notion of sustainability.

 

Troubling (Monolithic) Sustainability

 

In order to be able to unveil and critically analyze the propositions and suppositions of what I call ‘the monolithic sustainability discourse,’ it is fundamental to start with the etymology of the word ‘sustainability’ itself. The operationalization of the term can be problematic for it implies prior judgments about what is deemed important or necessary to sustain. While some of these judgements might resonate with an array of environmentalists who perceive that the health of the planet and the well-being of our descendants are being—or are already—compromised by certain human activities, various other perilous premises and assumptions are generally left unacknowledged as a result of the depoliticized character of the dominant discourse of sustainability. Lele and Norgaard (1996) have put forward three questions that can help us to uncover and think more critically about these presuppositions in and across various contexts and scales: (a) what is to be sustained, at what scale, and in what form?; (b) over what time period, with what level of certainty?; (c) through what social process(es), and with what trade-offs against other social goals? (p. 355).

 

By building on these critical questions and clarifications, we can better understand the nuances of how the destructive and thus unsustainable ethos of dehumanization and socio- ecological exploitation may inform and permeate normative notions and articulations of sustainability. Yet, this is only plausible if sustainability is politicized. To politicize is to engage the existing state of socio-political affairs, to problematize that which is taken for granted, to make explicit the power relations that are an innate part of everyday life and experience (Bailey & Gayle, 2003). In an attempt to comprehend why sustainability is typically depoliticized we ought to examine briefly its discursive history.

 

The term ‘sustainable development’ became a part of the policy discourse and almost every day language following the release of the Brundtland Commission’s report on the global environment and development in 1987 (Redclift, 2005). While their definition included a very clear social directive, its human and political dimensions have been largely overlooked amongst references to sustainability, which, due to its environmental origins (Lele & Norgaard, 1996) and neoliberal focus on rights rather than needs (Redclift, 2005), have typically focused on bio- physical, ecological issues (Vallance, Perkins, & Dixon, 2011). Social sustainability, which has been conceptualized in response to the failure of the sustainability approach to engender substantial change (Vallance et al., 2011), is the least developed of the three realms and is frequently framed in relation to ecological and/or economic sustainability (Magis & Shinn, 2013). I assert that the reason for this is twofold: first and foremost, the sustainability agenda was conceived by international committees and NGO networks, think tanks, and governmental structures (Agyeman et al., 2002), which makes it a top-down approach and, consequently, less likely to recognize and address themes such as structural poverty, equity, and justice (Colantonio, 2009); and second, because social sustainability is made subservient to economics and the environment, it fails to examine the socio-political circumstances and elements that are needed to sustain a community of people (Magis & Shinn, 2013).

 

Sustainability, since its inception as a Western construct, has been progressively viewed as a crucial driver in economic development and environmental management worldwide. Nevertheless, as delineated above, its almost universal focus on reconciling the growth model of economics and the environment has served to covertly depoliticize the dominant discourse and therefore render it uncontentious if not intrinsically benign. It is worth further exploring the dynamics of depoliticization for I believe they are at the radicle of the issues sustainability attempts to address in the first place.

 

Bailey and Gayle (2003) identify a series of acts that can be associated with the dynamics of depoliticization, three of which can be observed when examining the monolithic sustainability discourse: (a) eschewing political discourse; (b) removing from the discourse the recognition that social advantages are given to certain constituent groups; (c) not disclosing underlying viewpoints or values. These processes are enmeshed with intricate ideological instances that help to mask the systemic and/or structural nature of a social or cultural matter (Bailey & Gayle, 2003). Further, as Foucault (1984) has stated, “power is everywhere” (p. 93) and it is embodied and enacted in discourse and knowledge. Hence, possessing the analytical tools to name and unpack these discursive ideological formations and power dynamics ought to be a prerequisite to the development of more holistic and critically conscious understandings and applications of sustainability.

 

Politicizing Sustainability

 

If we are to envision and construe actual sustainable futures, we must first understand what brought us here, where the roots of the problems lie, and how the sustainability discourse and framework tackle—or fail to tackle—them. To do this is to politicize sustainability, to build a critical perspective of and about sustainability. It is an act of conscientização (or conscientization), to borrow Paulo Freire’s seminal term, of cultivating critical consciousness and conscience (Freire, 1993). It is a call for the necessity to highlight, problematize, and disrupt what I have termed ‘the ethos of unsustainability’ and its interrelated ideologies of dehumanization and exploitation. Ultimately, to embrace a stance that fails to scrutinize the sources of degradation and exploitation is to uphold the power relations that sustain oppressive structures (Freire, 1993; Perry, 2001). I assert that only by delving into the origins of the ‘ethos of unsustainability’ can we really devise sustainability paradigms that are capable of promoting significant socio-cultural transformation.

 

To comprehend the contours of the predicaments that loom on our horizon as well as their premises and logics, we must go back over 500 years in history to 1492, the year that marks the beginning of the current colonial era and the globalization of the European colonial imaginary (Tuck and Yang, 2012). It is important to note that my intention in doing so is not to provide a sweeping, all-encompassing description of this genealogy/historical process, but rather, to simply name, connect, and emphasize the ideological systems and patterns that have been conceptualized and reconceptualized so as to sustain the ethos of unsustainability and its exploitative power structures. After all, as Freire (1993) has indicated, “to name the world is to change it” (p. 88).

 

(World) Capitalism: A Technology of European Colonialism

 

According to the Oxford English Dictionary (OED), the word ‘colonialism’ stems from the Roman word ‘colonia,’ which meant ‘settlement’ or ‘farm.’ The OED describes it as:

 

… a body of people who settle in a new locality, forming a community subject to or connected with their parent state; the community so formed, consisting of the original settlers and their descendants and successors, as long as the connection with the parent state is kept up.

 

In Colonialism/Postcolonialism, Ania Loomba (2001) points out that this definition fails to link the word ‘colonialism’ to its ideologies of conquest and domination as it eschews any testimonial about those peoples who were already living in the places where the colonies were formalized. She offers another, more nuanced definition that hints to the processes of conquest and control of other peoples’ land and resources (Loomba, 2001, p. 2):

 

The process of ‘forming a community’ in the new land necessarily meant unforming or re-forming the communities that existed there already, and involved a wide range of practices including trade, plunder, negotiation, warfare, genocide, slavement and rebellions.

 

Loomba (2001) illuminates that while European colonialisms from the late fifteenth century onwards included a miscellany of patterns of domination and exploitation, it was a combination of these patterns that generated the economic disparity required for the maturation and expansion of European capitalism and industrial civilization; thus, capitalism demands the maintenance of colonial expansion in order to flourish. In spite of colonialism not being a monopoly of capitalism because it could be—and has been—utilized by so-called ‘socialist’ or ‘communist’ states as well (Dirlik, 2002), capitalism is a technology of colonialism that has been developed and re-structured over time as a means of advancing European colonial projects (Tuck and Yang, 2012). Colonialism was the instrument through which capitalism was able to reach its status as a global, master frame (Loomba, 2001).

 

A distinction between the three historical modes of colonialism might help to further elucidate the interrelationships between capitalism and colonialism.

 

Theories of coloniality as well as postcolonial theories typically acknowledge two brands of colonialism: external colonialism, which involves the appropriation of elements of Indigenous worlds in order to build the wealth and the power of the colonizers—the first world—, and internal colonialism, the bio- and geo-political management of people and land within the borders of a particular nation-state (Tuck and Yang, 2012). A third form, settler colonialism, is more suitable to describe the operationalization of colonialisms in which the colonizers arrive and make a new home on the land (Tuck and Yang, 2012). The settler objective of gaining control over land and resources by removing the local, Indigenous communities is an ongoing structure that relies on private property schemes and coercive systems of labor (Glenn, 2015).

 

In these processes of colonialism, land is conceived primarily if not exclusively as commodity and property, and human relationships to the land are only legitimized in terms of economic ownership (Tuck and Yang, 2012). These combined colonialist ideologies of commodification and private property are at the core of the various political economies of capitalism that are found in today’s globalized world (O’Sullivan, 2005). By relying on the appropriation of land and commodities through the “elimination of the Native” (Wolfe, 2006, p. 387), European colonialisms wind up restructuring non-capitalist economies so as to fuel European capitalism (Loomba, 2001). The globalization of the world is thereby the pinnacle of a process that started with the formation of the United States of America as the epitome of a Euro- centered, settler colonialist world power (Quijano, 2000).

 

Inspired by the European colonial imaginary, which transforms differences and diversity into a hierarchy of values (Mignolo, 2000) as well as by economic liberalism, which erases the production and labor contexts from the economy (Straume, 2011), the capitalist imaginary constitutes a broad depoliticization that disconnects its ‘social imaginary significations’ from the political sphere (Straume, 2011). Given that capitalism is imbued with European diffusionist constructs (Blaut, 1989), namely ‘progress,’ ‘development,’ and ‘modernity,’ the depoliticization of this now globalized imaginary is required not only to maintain the resilience of capitalism as a master frame (Straume, 2011), but also to camouflage its interconnectedness to European colonial systems.

 

Antonio Gramsci’s (1971) study and articulation of the conceptualization and operation of ideologies proves fruitful in terms of understanding how the capitalist imaginary has been used to facilitate processes of globalization that benefit European colonialisms. He argued that ideologies are invaluable when manufacturing consent as they are the means through which certain ideas and meanings are not only transmitted, but held to be true (Gramsci, 1971). Hence, hegemony, the power garnered through a combination of ideologies and coercion, is attained by playing with people’s common sense (Gramsci, 1971) and their lived system of meanings and values (Williams, 1976; see Loomba, 2001). Since subjectivity and ideology are key to the expansionist capitalist endeavor and its interrelated logics of commodification and domination (Gramsci, 1971), it becomes necessary to summon and dissect the colonial ideas and belief systems that have served and continue to serve as its conduits. This can in turn help us to interrogate the value systems and mental models that directly and/or indirectly inform the dominant notion of sustainability (education).

 

White Supremacist, Heteropatriarchal State Capitalism

 

As devised and practiced by Europeans and, later, by other Euro-centered powers such as the United States, colonial ideologies of race and racial structures smooth the way for capitalist production (Wolfe, 2006). The Eurocentric construct of race as “a system of discrimination, hierarchy and power” (Olson, 2004, xvii, p. 127-128) conveys colonial experience and infuses the most essential realms of world power and its hierarchies (Quijano, 2000). The state and its many institutions are particularly pivotal in sustaining these racialized ideologies that are obligatory for the development and continuance of capitalism (Loomba, 2001).

 

Slavery, as the foundation of notions of race and capitalist empire and one of the pillars of white supremacy, marks the concepts of ‘progress’ and ‘development’ as white (Painter, 2010) and renders black people as innately enslaveable, as nothing more than private property (Smith, 2010a). Within the context of the United States, the forms of slavery can and, indeed, have changed—from chattel slavery, to sharecropping, and more recently, to the prison industrial complex, which is still grounded in the premise that black bodies are an indefinite property of the state (Smith, 2010a)—yet, slavery as a logic of white supremacy has persisted (Smith, 2010a). The other two pillars of white supremacy are genocide, which expresses the need for Indigenous Peoples to always be disappearing, and orientalism, which builds on Edward Said’s influential term to explain how certain peoples and/or nations are coded as inferior and, therefore, a constant threat to the security and longevity of imperial states (Smith, 2010a).

 

The pillars of white supremacy may vary according to historical and geographical contexts (Smith, 2010a). Nonetheless, the centering of whiteness is generally what defines a colonial project. The formation of whiteness, or white identity, as a racialized class orientation stems from political efforts by capitalist elites and lawmakers to divide and conquer large masses of workers (Battalora, 2013). White identity is perhaps one of the most successful colonial and capitalist inventions since it “operates as a kind of property … with effects on social confidence and performance that can be empirically documented” (Alcoff, 2015, p. 23). It is a very dynamic category that can be enlarged to extend its privileges to others when white supremacist social and economic relations are jeopardized (Painter, 2010). It sustains itself, at least partially, by evading scrutiny and shifting the discursive focus to ‘non-whites’ (Silva, 2007). Whiteness is to be made invisible by remaining the norm, the standard, that which ought not to be questioned.

 

Capitalism therefore depends on and magnifies these racial hierarchies centered on whiteness. And, since race is imbricated and constructed simultaneously with gender, sexuality, ability, and other colonial categories—a conceptualization that serves to obscure white supremacy in state discourses and interventions (Kandaswamy, 2012)—, it is crucial to investigate the other ideologies that also shape class formation processes.

 

Heteropatriarchy, the combination of patriarchal and heterosexual control based on rigid and dichotomous gender identities—man and woman—and sexual orientations—heterosexual and homosexual—where one identity or orientation dominates the other, is another building block of colonialism. Patriarchy is employed to naturalize hierarchical relations within families and at a larger, societal level (Smith, 2010b). Similarly, heteronormativity paints heterosexual nuclear-domestic arrangements as normative (Arvin, Tuck, and Morrill, 2013) and is thus the bedrock of the colonial nation-state (Smith, 2010b). These social and cultural systems that configure heteropatriarchy are then apprehended as normal and natural whereas other arrangements or proclivities are demonized and perceived as repulsive and abnormal (Arvin et al., 2013). Heteropatriarchy is directly linked to colonial racial relations as it portrays white manhood as supreme and entitled to control over private property and to political sovereignty (Glenn, 2015). This indicates that the process of producing and managing gender frequently functions as a racial project that normalizes whiteness (Kandaswamy, 2012).

 

The laws and policies that were designed to institutionalize the formation of whiteness and white supremacy demonstrate that race, class, and gender are intertwined systems that uphold, constitute, and reconstitute each other (Battalora, 2013). The state and its ideological institutions are therefore major sites of racial struggle (Kandaswamy, 2012); they are responsible for devising and constantly revising the rationale that guides a white supremacist, heteropatriarchal settler colonialism grounded in the need to manufacture collective consent. These discourses are rooted in a pervasive state process that combines coercive state arbitration with societal consent by articulating the ideologies that link racial structure and representation as an effort to reorganize and distribute resources according to specific racial lines (Ferguson, 2012).

 

Despite increasing globalizing neoliberal urges toward deregulation and privatization, capitalism is still enabled and supported by the state. Its ‘ideological apparatuses,’ the state institutions and ideologies that enable and support the classist structure of capitalist societies (Althusser, 1989), is still fundamental to the expansion of capitalist enterprises; the nation-state is capitalism’s atomic component. The neoliberal state has utilized innovations in methods of social discipline and control along with legal practices to facilitate the process of economic globalization (Gill, 1995). Yet, all these schemes that involve retention of power through dominance and manufactured consent are rooted in divide and conquer strategies that cause those in subservient positions in society to engage in conflicts with one another (Hagopian, 2015). The interlinked logics and ideologies of white supremacy and heteropatriarchy conceived by state capitalism serve to spur dissent between potential opponents and thereby further stratify socio-economic classes. This prevents them from building a unified basis that can present a tangible threat to the status quo (Hagopian, 2015). Colonial and neocolonial powers have repeatedly deployed this stratagem to not only increase their geographical reach, but also to normalize and standardize the economic growth model of capitalism.

 

Colonialism is hence not just an ancient, bygone incident. The ideologies and processes delineated above demonstrate that it has remained very much in effect within contemporary capitalist and neoliberal frameworks (Preston, 2013). It then becomes critical to investigate how the dominant sustainability discourse may or may not collude in these schemes so that we may conceive of holistic blueprints that beget positive socio-ecological transformation.

 

Sustainability and Colonialism: Contradiction or Conscious Ideological Maneuver?

 

By unearthing what I believe are the roots of the predicament that sustainability attempts to heal, namely the ethos of dehumanization and exploitation rooted in divide and conquer systems, it becomes easier to analyze how the colonial political economy of capitalism may conserve hegemonic ideologies that pervade social relations and knowledge generating processes.

 

Yet, these ideologies and knowledge schemes have been given minimal attention in sustainability (education) scholarship. Even though some academics have contributed to the generation of a more critical comprehension of the interrelationships between capitalism, environmental degradation, and socio-economic justice (see Cachelin, Rose, & Paisley, 2015; Martusewicz, Edmundson, & Lupinacci, 2011; Pellow & Brulle, 2005), this major blindspot in linking sustainability to the colonial imaginary and its legacies prompts the following questions:(awhy are critiques of colonialism and capitalism so infrequent in the sustainability literature?: (a) why are critiques of colonialism and capitalism so infrequent in the sustainability literature?; and (b) how does that impact the discourse of sustainability?

 

I assert that, in spite of calls for paradigm shifts, the dominant disancourse of sustainability in the West embodies a transnational, globalized standard of economic growth. The promise that economic development can eradicate or at least alleviate poverty and hunger in a sustainable way reflects some of the same goals and values of the optimistic ‘ecological modernization’ concept and perspective, which suggest that the development and modernization of liberal capitalism result in improvements in ecological outcomes (Buttel, 2000). The neoliberal, capitalist overtones of sustainable development not only expose the contradiction inherent in the term, but they also serve to further commodify nature (Cock, 2011). This neoliberalization of nature, which has recently gained a lot of attention in the corporate world and academia under the lexicon of ‘ecosystem services,’ alienates people from their physical surroundings and therefore reinforces the society-nature divide. In short, the sustainability discourse has been appropriated by the capitalist master frame and has transformed most if not all social and ecological relations into financial ones. In lieu of addressing social and environmental justice issues, this form of “green” or “natural” capitalism is responsible for deepening both social and environmental inequalities (Cock, 2011).

 

Since sustainability (education) is (supposed to be) a praxis-oriented framework that symbiotically combines thought and action for transformative, liberatory ends, it ought to embrace this critique of colonial capitalism and the subsequent neoliberalization of the political economy if it is to oppose and resist hegemonic ideologies in its multiple and diverse manifestations. After all, whether intentionally or not, what matters in the end is that those discourses of sustainability that do not take a stance against colonialism and capitalism only serve to preserve them and the status quo. An understanding of these interdependent systems allows for the development of critical sustainability dialogues and actions that can actually promote the paradigmatic shifts required to redress the socio-cultural problems that are at the heart of the environmental crises. Thus, sustainability can and should be reframed to suggest a process of personal, social, and cultural conscientization that is environmentally sound, i.e. one that follows ecological principles and patterns, instead of upholding the dehumanizing, exploitative, and paradoxical ‘development as growth’ standard of global capitalism.

 

The following section combines the analyses and critiques presented in the preceding (sub)sections into a single, cohesive, and holistic framework, and further elucidates the distinctions between monolithic sustainability and critical sustainabilities.

 

The Framework of Critical Sustainability Studies

 

[T]he political cannot be restricted to a certain type of institution, or envisioned as constituting a specific sphere or level of society. It must be conceived as a dimension that is inherent to every human society and that determines our very ontological condition.

 

- Chantal Mouffe, The Return of the Political, 2005

 

‘Critical sustainability studies,’ while not exactly novel in the sense that it draws on principles, concepts, and positions that are foundational to other frameworks and fields—more specifically, critical Indigenous and ethnic studies, postcolonial theory, queer theory, feminist theory, crip theory, social ecology, political ecology, and cultural studies—, presents itself as an alternative to the sustainability theories and conceptualizations that have failed to engage a truly intersectional analysis of dominant sustainability and environmental discourses, policies, and practices. Its primary objective is to rearticulate sustainability as it has the potential to provide a more holistic conception of conscientization that can bridge the gap between social and economic justice and environmental sustainability.

 

The framework indicates a crucial double political intervention: to put sustainability and critical theory in conversation; to embed sustainability and ecology into critical theory and vice- versa. As I discussed in the previous section, sustainability has, for the most part, become a hegemonic and, therefore, highly problematic discourse that refuses to transform the complex ideologies and systems that undergird the ethos of unsustainability and the current socio- ecological crises. On the other hand, critical theory, which seeks to extend the consciousness of the human self as a social being within the context of dominant power structures and their knowledge management operations (Kincheloe, 2005), could benefit from incorporating ecological principles and the sustainability notion of ‘place’ into its analytical toolbox. After all, I am as interested in localizing critical knowledge—without disconnecting it from global matters and realities—as I am in putting forth more critical and radical views of sustainability. Hence, this framework brings together what I believe are some of the most robust and cutting edge theories and methodologies to facilitate the deconstruction of the questionable ideologies that guide Western epistemologies like (hegemonic) sustainability.

 

Critical sustainability studies encourages sustainability scholars and/or educators to move from a defined methodology of problem-solving to the more critical moment of calling something into question (Freire, 1993). By rooting it in conscientization, I propose an orientation to sustainability and sustainable development that politicizes and reveals it as an agenda, discourse, and knowledge system that ought to be contested and rearticulated so that it can incorporate and critically engage with emancipatory understandings of power and power relations. Furthermore, by problematizing and closing the culture-nature divide, it can lay down the groundwork for the paradigmatic changes necessary to heal widespread colonialist alienation from the wider ecological community and to create visions of deep sustainabilities that can engender ecologically sound socio-cultural transformation.

 

I stress that the notion of sustainabilities is necessary if we have the intention of opposing and displacing the monolithic, top-down and now universalized sustainability agenda, which I refer to as ‘big S Sustainability.’ After all, much like science (Parry, 2006), sustainability is not the property of any one culture or language. There are different ways of seeing and knowing sustainability, so it is time to pluralize it in the literature and discourse. This simple act is an extraordinary intervention in itself because within the colonial imaginary “sustainability” means “Western sustainability.” By centering “novel” understandings of sustainability that are concerned with the specificities of geo-political, cultural, and historical contexts and power relations, sustainability scholars and educators can create theories and visions of sustainability that can lead to the development of more just, place-based cultures and social ecologies.

 

Critical sustainability studies as I envision it is a consciousness-raising exercise that is particularly useful in educational settings. It indicates methodology as much as content. This praxis-oriented framework can help teachers and students alike to develop consciousness of freedom and to acknowledge authoritarian socio-cultural tendencies that have toxic environmental ramifications. The next section provides an overview of its tenets, the educational philosophy that underpins it, as well as the four preliminary methodological principles and examples of related pedagogical interventions that directly inform the framework and its liberatory, decolonizing ambitions.

 

Epistemological Position, Preliminary Methodological Principles, and Pedagogical Interventions for Conscientization

 

The epistemological, methodological, and pedagogical implications of critical sustainability studies are rooted in an ethical and political vision, one that is found in the vast majority of social ecology and political ecology projects: that “the domination of nature by man [sic] stems from the very real domination of human by human” (Bookchin, 2005, p. 1). In other words, we cannot overcome the ecological crisis unless we rid ourselves of the colonial ideologies of domination and hierarchy that permeate all forms of systemic and systematic exploitation and dehumanization. While much easier said than done, critical sustainability studies seeks to conceptualize this vision by building on the following tenets:

 

That sustainability and sustainability education are not neutral, they either advance or regress justice and Critical sustainability studies strives to promote justice and ecological regeneration.

That an analysis of power is central to understanding and engendering positive socio-cultural Critical sustainability studies strives to be conscious of power relations and to identify power inequalities and their implications.

That it is crucial to foreground the sociocultural identities and experiences of those who have been (most) oppressed – people of color, people with disabilities, queer and transgender people, the working class and the economically poor, undocumented immigrants, Critical sustainability studies acknowledges that just, healthy cultures and societies can only be cultivated if we examine the circumstances that cause and maintain socio-economic marginalization.

That positive socio-cultural transformation comes from the bottom up. Critical sustainability studies emphasizes and advocates a collective and decentralized approach to sustainable change.

And, finally, that the human community is inherently a part of rather than apart from the wider ecological world. Critical sustainability studies affirms that this relational ethos serves as the epistemological foundation of novel, dynamic worlds where healing and justice are at the front and center of our cultural and ecological identities.

In addition to delineating critical sustainability studies as a praxis that is founded on the above tenets, the framework is guided by a critical constructivist epistemological position. Strongly influenced by Freirean pedagogies and the Frankfurt school of thought, critical constructivism endeavors to dissect the processes by which knowledge is socially constructed; in other words, what we know about the worlds we live in always demands a knower and that which is to be known, a contextual and dialectical process that informs what we conceive of as reality (Kincheloe, 2005). This epistemological position problematizes and extends constructivism by illuminating the need for both teachers and students to develop a critical awareness of self, their perspectives, and ways their consciousness have been shaped and/or reshaped by society (Watts, Jofili, & Bezerra, 1997). Critical constructivists attempt to comprehend the forces that construe consciousness and the ways of seeing and being of the subjects who inhabit it (Kincheloe, 1993, as cited in Watts et al., 1997). This political, counter- Cartesianism, and anti-objectivist philosophy (Kincheloe, 2005) is central to an emancipatory approach to sustainability and sustainability education, and is, therefore, at the root of the critical sustainability studies conception of holistic conscientization.

 

www.susted.com/wordpress/content/critical-sustainability-...

Can you spot the similarities?

...The hairstyle

...The side cutouts

As Robyne said, they have a "Yin Yang" concept to them.

 

In Chinese philosophy, yin and yang is a concept of dualism in ancient Chinese philosophy, describing how seemingly opposite or contrary forces may actually be complementary, interconnected, and interdependent in the natural world, and how they may give rise to each other as they interrelate to one another.

Everglades National Park is an American national park that protects the southern twenty percent of the original Everglades in Florida. The park is the largest tropical wilderness in the United States and the largest wilderness of any kind east of the Mississippi River. An average of one million people visit the park each year. Everglades is the third-largest national park in the contiguous United States after Death Valley and Yellowstone. UNESCO declared the Everglades & Dry Tortugas Biosphere Reserve in 1976 and listed the park as a World Heritage Site in 1979, and the Ramsar Convention included the park on its list of Wetlands of International Importance in 1987. Everglades is one of only three locations in the world to appear on all three lists.

 

Most national parks preserve unique geographic features; Everglades National Park was the first created to protect a fragile ecosystem. The Everglades are a network of wetlands and forests fed by a river flowing 0.25 miles (0.40 km) per day out of Lake Okeechobee, southwest into Florida Bay. The park is the most significant breeding ground for tropical wading birds in North America and contains the largest mangrove ecosystem in the Western Hemisphere. Thirty-six threatened or protected species inhabit the park, including the Florida panther, the American crocodile, and the West Indian manatee, along with 350 species of birds, 300 species of fresh and saltwater fish, 40 species of mammals, and 50 species of reptiles. The majority of South Florida's fresh water, which is stored in the Biscayne Aquifer, is recharged in the park.

 

Humans have lived for thousands of years in or around the Everglades. Plans arose in 1882 to drain the wetlands and develop the land for agricultural and residential use. As the 20th century progressed, water flow from Lake Okeechobee was increasingly controlled and diverted to enable explosive growth of the Miami metropolitan area. The park was established in 1934, to protect the quickly vanishing Everglades, and dedicated in 1947, as major canal-building projects were initiated across South Florida. The ecosystems in Everglades National Park have suffered significantly from human activity, and restoration of the Everglades is a politically charged issue in South Florida.

 

Everglades National Park covers 1,508,976 acres (2,357.8 sq mi; 6,106.6 km2), throughout Dade, Monroe, and Collier counties in Florida, at the southern tip of the Atlantic coastal plain. The elevation typically ranges from 0 to 8 feet (2.4 m) above sea level, but a Calusa-built shell mound on the Gulf Coast rises 20 feet (6.1 m) above sea level.

 

The terrain of South Florida is relatively and consistently flat. The limestone that underlies the Everglades is integral to the diverse ecosystems within the park. Florida was once part of the African portion of the supercontinent Gondwana. After it separated, conditions allowed a shallow marine environment to deposit calcium carbonate in sand, shells, and coral to be converted into limestone. Tiny bits of shell, sand, and bryozoans compressed over multiple layers forming structures in the limestone called ooids, which created permeable conditions that hold water.

 

The Florida peninsula appeared above sea level between 100,000 and 150,000 years ago. As sea levels rose at the end of the Wisconsin ice age, the water table appeared closer to land. Lake Okeechobee began to flood, and convection thunderstorms were created. Vast peat deposits south of Lake Okeechobee indicate that regular flooding had occurred about 5,000 years ago. Plants began to migrate, subtropical ones from the northern part of Florida, and tropicals carried as seeds by birds from islands in the Caribbean. The limestone shelf appears to be flat, but there are slight rises—called pinnacles—and depressions caused by the erosion of limestone by the acidic properties of the water. The amount of time throughout the year that water is present in a location in the Everglades determines the type of soil, of which there only two in the Everglades: peat, created by many years of decomposing plant matter, and marl, the result of dried periphyton, or chunks of algae and microorganisms that create a grayish mud. Portions of the Everglades that remain flooded for more than nine months out of the year are usually covered by peat. Areas that are flooded for six months or less are covered by marl. Plant communities are determined by the type of soil and the amount of water present.

 

According to the Köppen climate classification system, Royal Palm at Everglades National Park has a tropical monsoon climate (Am). Summers are long, hot, and very wet and winters are warm and dry.

 

While they are common in the northern portion of Florida, no underground springs feed water into the Everglades system. An underground reservoir called the Floridan aquifer lies about 1,000 feet (300 m) below the surface of South Florida. The Everglades has an immense capacity for water storage, owing to the permeable limestone beneath the exposed land. Most of the water arrives in the form of rainfall, and a significant amount is stored in the limestone. Water evaporating from the Everglades becomes rain over metropolitan areas, providing the fresh water supply for the region. Water also flows into the park after falling as rain to the north onto the watersheds of the Kissimmee River and other sources of Lake Okeechobee, to appear in the Everglades days later. Water overflows Lake Okeechobee into a river 40 to 70 miles (64 to 113 km) wide, which moves almost imperceptibly.

 

At the turn of the 20th century, common concepts of what should be protected in national parks invariably included formidable geologic features like mountains, geysers, or canyons. As Florida's population began to grow significantly and urban areas near the Everglades were developed, proponents of the park's establishment faced difficulty in persuading the federal government and the people of Florida that the subtle and constantly shifting ecosystems in the Everglades were just as worthy of protection. When the park was established in 1947, it became the first area within the U.S. to protect flora and fauna native to a region as opposed to geologic scenery. The National Park Service recognizes nine distinct interdependent ecosystems within the park that constantly shift in size owing to the amount of water present and other environmental factors.

 

Freshwater sloughs are perhaps the most common ecosystem associated with Everglades National Park. These drainage channels are characterized by low-lying areas covered in fresh water, flowing at an almost imperceptible 100 feet (30 m) per day. Shark River Slough and Taylor Slough are significant features of the park. Sawgrass growing to a height of 6 feet (1.8 m) or more, and broad-leafed marsh plants, are so prominent in this region that they gave the Everglades its nickname "River of Grass", cemented in the public imagination in the title for Marjory Stoneman Douglas's book (1947), which culminated years of her advocacy for considering the Everglades ecosystem as more than a "swamp". Excellent feeding locations for birds, sloughs in the Everglades attract a great variety of waders such as herons, egrets, roseate spoonbills (Platalea ajaja), ibises and brown pelicans (Pelecanus occidentalis), as well as limpkins (Aramus guarauna) and snail kites that eat apple snails, which in turn feed on the sawgrass. The sloughs' availability of fish, amphibians, and young birds attract a variety of freshwater turtles, alligator (Alligator mississippiensis), water moccasin (Agkistrodon piscivorus conanti), and eastern diamondback rattlesnake (Crotalus adamanteus).

 

Freshwater marl prairies are similar to sloughs but lack the slow movement of surface water; instead, water seeps through a calcitic mud called marl. Algae and other microscopic organisms form periphyton, which attaches to limestone. When it dries it turns into a gray mud. Sawgrass and other water plants grow shorter in freshwater marl than they do in peat, the other type of soil in the Everglades which is found where water remains present longer throughout the year. Marl prairies are usually under water from three to seven months of the year, whereas sloughs may remain submerged for longer than nine months and sometimes remain under water from one year to the next. Sawgrass may dominate sloughs, creating a monoculture. Other grasses, such as muhly grass (Muhlenbergia sericea) and broad-leafed water plants can be found in marl prairies. Animals living in the freshwater sloughs also inhabit marl prairies. Marl prairies may go dry in some parts of the year; alligators play a vital role in maintaining life in remote parts of the Everglades by burrowing in the mud during the dry season, creating pools of water where fish and amphibians survive from one year to the next. Alligator holes also attract other animals who congregate to feed on smaller prey. When the region floods again during the wet season, the fish and amphibians which were sustained in the alligator holes then repopulate freshwater marl prairies.

 

Hammocks are often the only dry land within the park. They rise several inches above the grass-covered river and are dominated by diverse plant life consisting of subtropical and tropical trees, such as large southern live oaks (Quercus virginiana). Trees often form canopies under which animals thrive amongst scrub bushes of wild coffee (Psychotria), white indigoberry (Randia aculeata), poisonwood (Metopium toxiferum) and saw palmetto (Serenoa repens). The park features thousands of these tree islands amid sloughs—which often form the shape of a teardrop when seen from above (see park map) because of the slowly moving water around them—but they can also be found in pineland and mangroves. Trees in the Everglades, including wild tamarind (Lysiloma latisiliquum) and gumbo-limbo (Bursera simaruba), rarely grow higher than 50 feet (15 m) because of wind, fire, and climate.

 

The plant growth around the hammock base is nearly impenetrable; beneath the canopy hammocks is an ideal habitat for animals. Reptiles (such as various species of snake and anole) and amphibians (such as the American green tree frog, Hyla cinerea), live in the hardwood hammocks. Birds such as barred owls (Strix varia), woodpeckers, northern cardinals (Cardinalis cardinalis), and southern bald eagles (Haliaeetus leucocephalus leucocephalus) nest in hammock trees. Mammal species living in hardwood hammocks include Florida black bears (Ursus americanus floridanus), red foxes (Vulpes vulpes), minks (Neogale vison), marsh rabbits (Sylvilagus palustris), gray foxes (Urocyon cinereoargenteus), white-tailed deer (Odocoileus virginianus), and the rare, critically endangered Florida panther (Puma concolor couguar).

 

Miami-Dade County was once covered in 186,000 acres (290.6 sq mi; 752.7 km2) of pine rockland forests, but most of it was harvested by the lumber industry. Pineland ecosystems (or pine rocklands) are characterized by shallow, dry sandy loam over a limestone substrate covered almost exclusively by slash pines (Pinus elliottii var. densa). Trees in this ecosystem grow in solution holes, where the soft limestone has worn away and filled with soil, allowing plants to take hold. Pinelands require regular maintenance by fire to ensure their existence. South Florida slash pines are uniquely adapted to promote fire by dropping a large amount of dried pine needles and shedding dry bark. Pine cones require heat from fires to open, allowing seeds to disperse and take hold. The trunks and roots of slash pines are resistant to fire. Prescribed burns in these areas take place every three to seven years; without regular fires, hardwood trees begin to grow in this region, and pinelands become recategorized as mixed swamp forests. Most plants in the area bloom about 16 weeks after a fire. Nearly all pinelands have an understory of palm shrubs and a diverse ground covering of wild herbs.

 

Pine rocklands are considered one of the most threatened habitats in Florida; less than 4,000 acres (6.3 sq mi; 16.2 km2) of pineland exist outside the park. Within the park, 20,000 acres (31.3 sq mi; 80.9 km2) of pineland are protected. A variety of animal species meet their needs for food, shelter, nesting, and rooking in pine rocklands. Woodpeckers, eastern meadowlarks (Sturnella magna), loggerhead shrikes (Lanius ludovicianus), grackles, and northern mockingbirds (Mimus polyglottos) are commonly found in pinelands. Black bears and Florida panthers also live in this habitat.

 

Cypress trees are conifers that are adapted to live in standing fresh water. They grow in compact structures called cypress domes and in long strands over limestone. Water levels may fluctuate dramatically around cypress domes and strands, so cypresses develop "knees" that protrude from the water at high levels to provide oxygen for the root systems. Dwarf cypress trees grow in drier areas with poorer soil. Epiphytes, such as bromeliads, Spanish moss (Tillandsia usneoides), orchids and ferns grow on the branches and trunks of cypress trees. Everglades National Park features twenty-five species of orchids. Tall cypress trees provide excellent nesting areas for birds including wild turkey (Meleagris gallopavo), ibis, herons, egrets, anhingas (Anhinga anhinga), and belted kingfisher (Megaceryle alcyon). Mammals in cypress regions include white-tailed deer, squirrels, raccoons, opossums, skunks, swamp rabbits, river otters (Lontra canadensis), and bobcats, as well as small rodents.

 

Mangrove trees cover the coastlines of South Florida, sometimes growing inland depending on the amount of salt water present within the Everglades ecosystems. During drier years when less fresh water flows to the coast, mangroves will appear among fresh water plants. When rain is abundant, sawgrass and other fresh water plants may be found closer to the coast. Three species of mangrove trees—red (Rhizophora mangle), black (Avicennia germinans), and white (Laguncularia racemosa)—can be found in the Everglades. With a high tolerance of salt water, winds, extreme tides, high temperatures, and muddy soils, mangrove trees are uniquely adapted to extreme conditions. They act as nurseries for many marine and bird species. They are also Florida's first defense against the destructive forces of hurricanes, absorbing flood waters and preventing coastal erosion. The mangrove system in Everglades National Park is the largest continuous system of mangroves in the world.

 

Within the Florida mangrove systems live 220 species of fish, and a variety of crabs, crayfish, shrimp, mollusks, and other invertebrates, which serve as the main source of food for many birds. Dozens of bird species use mangroves as nurseries and food stores, including pelicans, grebes, tricolored herons (Egretta tricolor), gulls, terns, hawks and kites, and arboreal birds like mangrove cuckoos (Coccyzus minor), yellow warblers (Dendroica petechia), and white-crowned pigeons (Patagioenas leucocephala). The mangroves also support 24 species of amphibians and reptiles, and 18 species of mammals, including the endangered green turtle (Chelonia mydas), hawksbill turtle (Eretmochelys imbricata), and West Indian manatee (Trichechus manatus).

 

Coastal lowlands, or wet prairies, are salt water marshes that absorb marine water when it gets high or fresh water when rains are heavy. Floods occur during hurricane and tropical storm surges when ocean water can rise several feet over the land. Heavy wet seasons also cause floods when rain from the north flows into the Everglades. Few trees can survive in the conditions of this region, but plants—succulents like saltwort and glasswort—tolerate salt, brackish water, and desert conditions. Animal life in this zone is dependent upon the amount of water present, but commonly found animals include Cape Sable seaside sparrow (Ammodramus maritimus mirabilis), Everglades snail kite (Rostrhamus sociabilis), wood stork (Mycteria americana), eastern indigo snake (Drymarchon couperi), and small mammals such as rats, mice, and rabbits.

 

The largest body of water within the park is Florida Bay, which extends from the mangrove swamps of the mainland's southern tip to the Florida Keys. Over 800 square miles (2,100 km2) of marine ecosystem lies in this range. Coral, sponges, and seagrasses serve as shelter and food for crustaceans and mollusks, which in turn are the primary food source for larger marine animals. Sharks, stingrays, and barracudas also live in this ecosystem. Pelicans, shorebirds, terns, and black skimmers (Rynchops niger) are among the birds frequenting park shorelines. The bay also has its own resident population of bottlenose dolphin (Tursiops truncatus).

 

The bay's many basins are broken up by sandbanks that serve as plentiful recreational fishing grounds for snook (Centropomus undecimalis), redfish (Sciaenops ocellatus), spotted seatrout (Cynoscion nebulosus), tarpon (Megaflops atlanticus), bonefish (Albula vulpes), and permit (Trichinous falcatus),[48] as well as snapper (Lutjanus campechanus), bluegill (Lepomis macrochirus), and bass. Wading birds such as roseate spoonbills (Platalea ajaja), reddish egrets (Egretta rufescens), and great white herons (Ardea herodias occidentalis) have unique subpopulations that are largely restricted to Florida Bay. Other bird species include bald eagles, cormorants, and ospreys. Mammals along the shoreline include raccoons, opossums, bobcats, and fox squirrels.

 

Humans likely first inhabited the South Florida region 10,000 to 20,000 years ago. Two tribes of Native Americans developed on the peninsula's southern tip: the Tequesta lived on the eastern side and the Calusa, greater in numbers, on the western side. The Everglades served as a natural boundary between them. The Tequesta lived in a single large community near the mouth of the Miami River, while the Calusa lived in 30 villages. Both groups traveled through the Everglades but rarely lived within them, remaining mostly along the coast.

 

The diets of both groups consisted mostly of shellfish and fish, small mammals, game, and wild plants. Having access only to soft limestone, most of the tools fashioned by Native Americans in the region were made of shell, bone, wood, and animal teeth; shark teeth were used as cutting blades, and sharpened reeds became arrows and spears. Shell mounds still exist today within the park, giving archaeologists and anthropologists evidence of the raw materials available to the indigenous people for tool construction. Spanish explorers estimated the number of Tequesta at first contact to be around 800, and Calusa at 2,000; the National Park Service reports there were probably about 20,000 natives living in or near the Everglades when the Spanish established contact in the late 16th century. The Calusa lived in social strata and were able to create canals, earthworks, and shellworks. The Calusa were also able to resist Spanish attempts at conquest.

 

The Spanish had contact with these societies and established missions further north, near Lake Okeechobee. In the 18th century, invading Creeks incorporated the dwindling numbers of the Tequesta into their own. Neither the Tequesta nor Calusa tribe existed by 1800. Disease, warfare, and capture for slavery were the reasons for the eradication of both groups. The only evidence of their existence within the park boundaries is a series of shell mounds that were built by the Calusa.

 

In the early 19th century, Creeks, escaped African slaves, and other Indians from northern Florida displaced by the Creek War, formed the area's Seminole nation. After the end of the Seminole Wars in 1842, the Seminoles faced relocation to Indian territory near Oklahoma. A few hundred Seminole hunters and scouts settled within what is today Big Cypress National Preserve, to escape the forced emigration to the west. From 1859 to about 1930, the Seminoles and Miccosukee, a similar but linguistically unique tribe, lived in relative isolation, making their living by trading. In 1928, surveying and construction began on the Tamiami Trail, along the northern border of Everglades National Park. The road bisected the Everglades, introducing a steady, if small, traffic of white settlers into the Everglades.

 

Some members of the Miccosukee and Seminole tribes continue to live within park boundaries. Management of the park includes approval of new policies and procedures by tribal representatives "in such a manner that they do not conflict with the park purpose".

 

Following the end of the Seminole Wars, Americans began settling at isolated points along the coast in what is now the park, from the Ten Thousand Islands to Cape Sable. Communities developed on the two largest pieces of dry ground in the area, on Chokoloskee Island and at Flamingo on Cape Sable, both of which established post offices in the early 1890s. Chokoloskee Island is a shell mound, a midden built roughly 20 feet (6 m) high over thousands of years of occupation by the Calusa. The settlements in Chokoloskee and Flamingo served as trading centers for small populations of farmers, fishermen, and charcoal burners settled in the Ten Thousand Islands. Both settlements and the more isolated homesteads could only be reached by boat until well into the 20th century. Everglades City, on the mainland near Chokoloskee, enjoyed a brief period of prosperity when, beginning in 1920, it served as the headquarters for the construction of the Tamiami Trail. A dirt road from Florida City reached Flamingo in 1922, while a causeway finally connected Chokoloskee to the mainland's Everglades City in 1956.

 

After the park was established, private property in the Flamingo area was claimed by eminent domain, and the site was incorporated into the park as a visitor center.

 

Several attempts were made to drain and develop the Everglades in the 1880s. The first canals built in the Everglades did little harm to the ecosystem, as they were unable to drain much of it. Napoleon Bonaparte Broward based the majority of his 1904 campaign for governor on how drainage would create "The Empire of the Everglades". Broward ordered the drainage that took place between 1905 and 1910, and it was successful enough that land developers sold tracts for $30 per acre, settling the town of Davie, and developing regions in Lee and Dade counties. The canals also cleared water that made way for agricultural fields growing sugarcane.

 

In the 1920s, a population boom in South Florida created the Florida land boom, which was described by author Michael Grunwald as "insanity". Land was sold before any homes or structures were built on it and in some cases before any plans for construction were in place. New landowners, eager to make good on their investments, hastily constructed homes and small towns on recently drained land. Mangrove trees on the coasts were taken down for better views and replaced with shallow-rooted palm trees. The U.S. Army Corps of Engineers began construction on larger canals to control the rising waters in the Everglades. Nevertheless, Lake Okeechobee continued to rise and fall, the region was covered with rain, and city planners continued to battle the water. The 1926 Miami Hurricane caused Lake Okeechobee levees to fail; hundreds of people south of the lake drowned. Two years later, the 1928 Okeechobee Hurricane claimed 2,500 lives when Lake Okeechobee once again surged over its levees. Politicians who declared the Everglades uninhabitable were silenced when a four-story wall, the Herbert Hoover Dike, was built around Lake Okeechobee. This wall effectively cut off the water source from the Everglades.

 

Following the wall's construction, South Florida endured a drought severe enough to cause serious wildfires in 1939. The influx of humans had a detrimental effect on the plants and animals of the region when melaleuca trees (Melaleuca quinquenervia) were introduced to help with drainage, along with Australian pines brought in by developers as windbreaks. The region's timber was devastated for lumber supplies. Alligators, birds, frogs, and fish were hunted on a large scale. Entire rookeries of wading birds were shot to collect their plumes, which were used in women's hats in the early 20th century. The largest impact people had on the region was the diversion of water away from the Everglades. Canals were deepened and widened, and water levels fell dramatically, causing chaos in food webs. Salt water replaced fresh water in the canals, and by 1997 scientists noticed that salt water was seeping into the Biscayne Aquifer, South Florida's water source.

 

In the 1940s, Marjory Stoneman Douglas, a freelance writer and former reporter for The Miami Herald, began to research the Everglades for an assignment about the Miami River. She studied the land and water for five years and published The Everglades: River of Grass in 1947, describing the area in great detail, including a chapter on its disappearance. She wrote: "What had been a river of grass and sweet water that had given meaning and life and uniqueness to this enormous geography through centuries in which man had no place here was made, in one chaotic gesture of greed and ignorance and folly, a river of fire." The book has sold 500,000 copies since its publication, and Douglas's continued dedication to ecology conservation earned her the nicknames "Grand Dame of the Everglades", "Grandmother of the Everglades" and "the anti-Christ" for her singular focus at the expense of some political interests. She founded and served as president for an organization called Friends of the Everglades, initially intended to protest the construction of a proposed Big Cypress jetport in 1968. Successful in that confrontation, the organization has grown to over 4,000 members, committed to the preservation of the Everglades. She wrote and spoke about the importance of the Everglades until her death at age 108 in 1998.

 

Floridians hoping to preserve at least part of the Everglades began to express their concern over diminishing resources in the early 20th century. Royal Palm State Park was created in 1916 and protected Paradise Key; it included several trails and a visitor center several miles from Homestead. Miami-based naturalists first proposed that the area become a national park in 1923. Five years later, the Florida state legislature established the Tropical Everglades National Park Commission to study the formation of a protected area. The commission was led by Ernest F. Coe, a land developer turned conservationist, who was eventually nicknamed Father of Everglades National Park. Coe's original plan for the park included more than 2,000,000 acres (3,125.0 sq mi; 8,093.7 km2) including Key Largo and Big Cypress, and his unwillingness to compromise almost prevented the park's creation. Various other interests, including land developers and sport hunters, demanded that the size of the park be decreased.

 

The commission was also tasked with proposing a method to raise the money to purchase the land. The search coincided with the arrival of the Great Depression in the United States, and money for land purchase was scarce. The U.S. House of Representatives authorized the creation of the new national park on May 30, 1934, but the Act (HR 2837), which permanently reserved lands donated by public or private donation as wilderness, passed only with a rider that ensured no money would be allotted to the project for at least five years. Coe's passion and U.S. Senator Spessard Holland's politicking helped to fully establish the park, after Holland was able to negotiate 1,300,000 acres (2,031.2 sq mi; 5,260.9 km2) of the park, leaving out Big Cypress, Key Largo, the Turner River area, and a 22,000-acre (34.4 sq mi; 89.0 km2) tract of land called "The Hole in the Donut" that was too highly valued for agriculture. Miami Herald editor John Pennekamp was instrumental in pushing the Florida Legislature to raise $2 million to purchase the private land inside the park boundaries. It was dedicated by President Harry Truman on December 6, 1947, one month after Marjory Stoneman Douglas's book The Everglades: River of Grass was released. The same year, several tropical storms struck South Florida, prompting the construction of 1,400 miles (2,300 km) of canals, sending water unwanted by farmers and residents to the ocean.

 

The Central and Southern Florida Flood Control Project (C&SF) was authorized by Congress to construct more than one thousand miles of canals and flood control structures across South Florida. The C&SF, run by the U.S. Army Corps of Engineers, established an agricultural area directly south of Lake Okeechobee, and three water conservation areas, all bordered by canals that diverted excess water either to urban areas or into the Atlantic Ocean, Gulf of Mexico or Florida Bay. South of these manmade regions was Everglades National Park, which had been effectively cut off from its water supply. By the 1960s, the park was visibly suffering. The C&SF was directed to provide enough water to sustain the park; it did not follow through. A proposed airport that would have dire environmental effects on Everglades National Park became the center of a battle that helped to initiate the environmental movement into local and national politics. The airport proposal was eventually abandoned, and in 1972 a bill was introduced to curb development in South Florida and ensure the national park would receive the amount of water it needed. Efforts turned to repairing the damage wrought by decades of mismanagement: the Army Corps of Engineers changed its focus in 1990 from constructing dams and canals to constructing "purely environmental projects".

 

Regions originally included in Ernest Coe's vision for a national park were slowly added over the years to the park or incorporated into other protected areas: Biscayne National Park, Big Cypress National Preserve, John Pennekamp Coral Reef State Park on Key Largo, Ten Thousand Islands National Wildlife Refuge, and Florida Keys National Marine Sanctuary were all protected after the park's opening in 1947. Everglades National Park was designated an International Biosphere Reserve on October 26, 1976. On November 10, 1978, 1,296,500 acres (2,025.8 sq mi; 5,246.7 km2), about 86% of the park, was declared a wilderness area. It was renamed the Marjory Stoneman Douglas Wilderness in 1997. It was listed as a UNESCO World Heritage Site on October 24, 1979, and as a Wetland of International Importance on June 4, 1987. It was placed on the List of World Heritage in Danger from 1993 until 2007 and then again in 2010. The park was added again due to the continued degradation of the set causing significant indications of eutrophication (for example algal blooms) negatively impacting the marine life causing the US government to request UNESCO and IUCN for assistance in development.

 

President George H. W. Bush signed the Everglades National Park Protection and Expansion Act on December 13, 1989, that added 109,506 acres (171.1 sq mi; 443.2 km2) to the eastern side of the park, closed the park to airboats, directed the Department of the Army to restore water to improve the ecosystems within Everglades National Park, and "Direct(ed) the Secretary of the Interior to manage the Park in order to maintain the natural abundance, diversity, and ecological integrity of native plants and animals, as well as the behavior of native animals, as part of their ecosystem." Bush remarked in his statement when signing the act, "Through this legislation that river of grass may now be restored to its natural flow of water".

 

In 2000, Congress approved the Comprehensive Everglades Restoration Plan (CERP), a federal effort to restore the Everglades with the objectives of "restoration, preservation and protection of the south Florida ecosystem while providing for other water-related needs of the region", and claiming to be the largest environmental restoration in history. It was a controversial plan; detractors worried that it "relies on uncertain technologies, overlooks water quality, subsidizes damaging growth and delays its environmental benefits".[90] Supporters of the plan included the National Audubon Society, who were accused by Friends of the Everglades and the Biodiversity Legal Foundation of prioritizing agricultural and business interests.

 

The namesake of Anhinga Trail dries its feathers

CERP projects are designed to capture 1.7 billion US gallons (6,400,000 m3) of fresh water every day, store it in underground reservoirs, and release the water to areas within 16 counties in South Florida. Approximately 35,600 acres (55.6 sq mi; 144.1 km2) of man-made wetlands are to be constructed to confine contaminated water before it is released to the Everglades, and 240 miles (390 km) of canals that divert water away from the Everglades are to be destroyed. During the first five years of implementation, CERP was responsible for the purchase of 207,000 acres (323.4 sq mi; 837.7 km2) of land at a cost of $1 billion. The plan aims to spend $10.5 billion over 30 years, combining 50 different projects and giving them 5-year timelines.

 

Everglades National Park was directly hit by Hurricanes Katrina, Wilma, and Rita in 2005. Such storms are a natural part of the park's ecosystem; 1960's Hurricane Donna left nothing in the mangroves but "standing dead snags" several miles wide, but 30 years later the area had completely recovered. Predictably, what suffered the most in the park from the 2005 hurricanes were man-made structures. In 2009 the visitor center and lodge at Flamingo were irreparably damaged by 125 mph (201 km/h) winds and an 8 ft (2.4 m) storm surge; the lodge had been functioning for 50 years when it was torn down; nothing is slated to replace it.

 

Everglades National Park reported in 2005 a budget of over $28 million. Of that, $14.8 million was granted from the National Park Service and $13.5 million from various sources including CERP, donations, and other grants. The entry fee for private vehicles in 2021 is $30. Of the nearly one million visitors to Everglades National Park in 2006, more than 38,000 were overnight campers, paying $16 a night or $10 a night for backcountry permits. Visitors spent $2.6 million within the park and $48 million in local economies. More than 900 jobs were sustained or created within or by the park, and the park added value of $35 million to local economies.

 

Everglades National Park has had 19 superintendents since it was dedicated in 1947. The park's first superintendent, Daniel Beard (1947-1958), was also its longest-serving. After Superintendent Beard, Warren F. Hamilton served between 1958 and 1963, followed by Stanley C. Joseph (1963-1966), Roger W. Allin (1966-1968), John C. Raftery (1968-1970), Joseph Brown (1970-1971), Jack E. Stark (1971-1976), John M. Good (1976-1980), John M. Morehead (1980-1986), Marueen E. Finnerty (Acting Superintendent, 1986), Michael V. Finley (1986-1989), Robert L. Arnberger (Acting Superintendent, 1989), Robert S. Chandler (1989-1992), Dick Ring (1992-2000), Marueen E. Finnerty (2000-2003), Dan Kimball (2004-2014), Shawn Benge (Acting Superintendent, 2014), Bob Krumenaker (Acting Superintendent, 2014-2015), and finally Pedro Ramos, who was appointed in 2015 and continues to serve.

 

The park was placed into Administrative Region I in 1937, when the regions were first established. Region I was retitled the Southeast Region in 1962, which was restructured into the Southeast Area in 1995. The reorganized unified Interior regions put it in the new Region 2.

 

The busiest season for visitors is from December to March, when temperatures are lowest and mosquitoes are least active. The park features five visitor centers: on the Tamiami Trail (part of U.S. Route 41) directly west of Miami is the Shark Valley Visitor Center. A fifteen-mile (24 km) round trip path leads from this center to a two-story observation tower. Tram tours are available during the busy season. Closest to Homestead on State Road 9336 is the Ernest F. Coe Visitor Center, where a 38-mile (61 km) road begins, winding through pine rockland, cypress, freshwater marl prairie, coastal prairie, and mangrove ecosystems. Various hiking trails are accessible from the road, which runs to the Flamingo Visitor Center and marina, open and staffed during the busier time of the year. The Gulf Coast Visitor Center is closest to Everglades City on State Road 29 along the west coast. The Gulf Coast Visitor Center gives canoers access to the Wilderness Waterway, a 99-mile (160 km) canoe trail that extends to the Flamingo Visitor Center. The former Royal Palm State Park was the site of the first Everglades National Park visitor center and later became the Royal Palm Visitor Center within the park. The western coast of the park and the Ten Thousand Islands and the various key islands in Florida Bay are accessible only by boat.

 

Several walking trails in the park vary in hiking difficulty on Pine Island, where visitors can cross hardwood hammocks, pinelands, and freshwater sloughs. Starting at the Royal Palm Visitor Center, the Anhinga Trail is a half-mile self-guided tour through a sawgrass marsh where visitors can see alligators, marsh and wading birds, turtles, and bromeliads. Its proximity to Homestead and its accessibility make it one of the most visited sites in the park. The nearby Gumbo Limbo Trail is also self-guided, at half-mile long. It loops through a canopy of hardwood hammocks that include gumbo limbo (Bursera simaruba), royal palms (Roystonea), strangler figs (Ficus aurea), and a variety of epiphytes.

 

Twenty-eight miles (45 km) of trails start near the Long Pine Key campgrounds and wind through Long Pine Key, well-suited for offroad cycling through the pine rocklands in the Marjory Stoneman Douglas Wilderness Area. Two boardwalks allow visitors to walk through a cypress forest at Pa-Hay-O-Kee, which also features a two-story overlook, and another at Mahogany Hammock (referring to Swietenia mahagoni) that takes hikers through a dense forest in the middle of a freshwater marl prairie. Closer to Flamingo, more rugged trails take visitors through mangrove swamps, along Florida Bay. Christian Point Trail, Snake Bight Trail, Rowdy Bend Trail and Coastal Prairie Trail allow viewing of shorebirds and wading birds among the mangroves. Portions of the trails may be impassable depending on the time of year, because of mosquitoes and water levels. Ranger-led tours take place in the busier season only.

 

Camping is available year-round in Everglades National Park. Camping with some services is available at Long Pine Key, close to the Ernest F. Coe Visitor Center, where 108 sites are accessible by car. Near Flamingo, 234 campsites with some services are also available. Recreational vehicle camping is available at these sites, but not with all necessary services. Back-country permits are required for campsites along the Wilderness Waterway, Gulf Coast sites, and sites in the various keys. Several back-country sites are chickees; others are beach and ground sites.

 

Low-powered motorboats are allowed in the park; the majority of salt water areas are no-wake zones to protect manatees and other marine animals from harm. Jet skis, airboats, and other motorized personal watercraft are prohibited. Many trails allow kayaks and canoes. A state license is required for fishing. Fresh water licenses are not sold in the park, but a salt water license may be available. Swimming is not recommended within the park boundaries; water moccasins, snapping turtles (Chelydra serpentina), alligators, and crocodiles thrive in fresh water. Sharks, barracuda, and sharp dangerous coral are plentiful in salt water. Visibility is low in both salt water and fresh water areas.

 

Everglades National Park is an important part of the Great Florida Birding Trail. It has great biodiversity and many species of birds for bird watching and bird photography also.

 

Portions of Everglades National Park are ideal for dark sky observations in South Florida. The best viewing locations are in the remote southern and western areas of the Everglades, such as Flamingo and the Ten Thousand Islands. The Milky Way appears brightest when looking south, toward the least light-polluted areas.

 

Less than 50 percent of the Everglades which existed prior to drainage attempts remains intact today. Populations of wading birds dwindled 90 percent from their original numbers between the 1940s and 2000s. The diversion of water to South Florida's still-growing metropolitan areas is the Everglades National Park's number one threat. In the 1950s and 1960s, 1,400 miles (2,300 km) of canals and levees, 150 gates and spillways, and 16 pumping stations were constructed to direct water toward cities and away from the Everglades. Low levels of water leave fish vulnerable to reptiles and birds, and as sawgrass dries it can burn or die off, which in turn kills apple snails and other animals that wading birds feed upon. Populations of birds fluctuate; in 2009, the South Florida Water Management District claimed wading birds across South Florida increased by 335 percent. Following three years of increasing numbers, The Miami Herald reported in 2009 that populations of wading birds within the park decreased by 29 percent.

 

Cities along the west coast of Florida rely on desalinization for fresh water; the quantity demanded is too great for the land to provide. Nitrates in the underground water system and high levels of mercury also impact the quality of fresh water the park receives. In 1998, a Florida panther was found dead in Shark Water Slough, with levels of mercury high enough to kill a human. Increased occurrences of algal blooms and red tide in Biscayne Bay and Florida Bay have been traced to the amounts of controlled water released from Lake Okeechobee. The brochure given to visitors at Everglades National Park includes a statement that reads, "Freshwater flowing into the park is engineered. With the help of pumps, floodgates, and retention ponds along the park's boundary, the Everglades is presently on life support, alive but diminished."

 

A series of levees on the park's eastern border marks the line between urban and protected areas, but development into these areas threatens the park system. Florida still attracts nearly a thousand new residents every day,[119] and building residential, commercial and industrial zones near Everglades National Park stresses the water balance and ecosystems within the park. On the park's western border, Fort Myers, Naples, and Cape Coral are expanding, but no system of levees exists to mark that border. National Geographic rated both Everglades National Park and Big Cypress National Preserve the lowest-scoring parks in North America, at 32 out of 100. Their scoring system rated 55 parks by their sustainable tourism, destination quality, and park management. The experts who compiled the results justified the score by stating: "Encroachment by housing and retail development has thrown the precious ecosystem into a tailspin, and if humankind doesn't back off, there will be nothing left of one of this country's most amazing treasures".

 

Thirty-six federally protected animals live in the park, some of which face grave threats to their survival.

 

In the United States, the American crocodile's only habitat is within South Florida. They were once overhunted for their hides. They are protected today from hunting but are still threatened by habitat destruction and injury from vehicle collisions when crossing roads to reach waterways. About 2,000 crocodiles live in Florida, and there are roughly 100 nests in the Everglades and Biscayne National Parks. Crocodiles populations in South Florida have increased as has the number of alligators. Crocodiles were reclassified from "endangered" to "threatened" in the United States in 2007.

 

The Florida panther is one of the most endangered mammals on earth. About 230 live in the wild, primarily in the Everglades and the Big Cypress Swamp. The biggest threats to the panther include habitat destruction from human development, vehicle collisions, inbreeding due to their limited gene pool, parasites, diseases, and mercury poisoning.

 

Four Everglade species of sea turtle including the Atlantic green sea turtle, the Atlantic hawksbill, the Atlantic loggerhead (Caretta caretta), and the Atlantic ridley (Lepidochelys kempii) are endangered. Also, the leatherback sea turtle (Dermochelys coriacea) is threatened. Numbers are difficult to determine, since males and juveniles do not return to their birthplace; females lay eggs in the same location every year. Habitat loss, illegal poaching, and destructive fishing practices are the biggest threats to these animals.

 

The range of the Cape Sable seaside sparrow is restricted to Everglades National Park and the Big Cypress Swamp. In 1981 6,656 Cape Sable seaside sparrows were reported in park boundaries, but surveys over 10 years documented a decline to an estimated 2,624 birds by 2002. Attempts to return natural levels of water to the park have been controversial; Cape Sable seaside sparrows nest about a foot off the ground, and rising water levels may harm future populations, as well as threaten the locally endangered snail kite. The Everglades snail kite eats apple snails almost exclusively, and the Everglades is the only location in the United States where this bird of prey exists. There is some evidence that the population may be increasing, but the loss of habitat and food sources keep the estimated number of these birds at several hundred.

 

The West Indian manatee has been upgraded from endangered to threatened. Collisions with boats and habitat loss are still its biggest threats.

 

Fire naturally occurs after lightning storms but takes its heaviest toll when water levels are low. Hardwood hammock and cypress trees are susceptible to heavy damage from fire, and some may take decades to grow back. Peat built up over centuries in the marsh can cause fires to burn deep scars in the soil. In 2007, Fred Sklar of the South Florida Water Management District said: "An extreme drought can be viewed (as) almost as catastrophic as a volcano. It can reshape the entire landscape. It can take 1,000 years to produce two inches of peat, and you can lose those couple of inches in a week."

 

Rising sea levels caused by global warming are another threat to the future of the park. Since 1932, ocean levels at Key West have steadily risen over 0.7 feet (0.2 m), which could have disastrous consequences for land so close to the ocean. It is estimated that within 500 years freshwater habitats in the Everglades National Park will be obliterated by salt water, leaving only the northernmost portion of the Everglades. Cost estimates for raising or replacing the Tamiami Trail and Alligator Alley with bridges are in the hundreds of millions of dollars.

 

Through Trump Administration, The Florida Department of Transportation, and Everglades National Park, there are plans to execute and complete the Next Steps project to help fix these various water issues, along with other parts of the park. This completion plan was announced in September 2020, will begin November 2020, and should be done by the end of 2024.

 

The introduction of non-native species into South Florida is a considerable problem for the park. Many of the biological controls such as weather, disease, and consumers who naturally limit plants in their native environments do not exist in the Everglades, causing many to grow larger and multiply far beyond their average numbers in their native habitats. Approximately 26 percent of all fish, reptiles, birds, and mammal species in South Florida are exotic—more than in any other part of the U.S.—and the region hosts one of the highest numbers of exotic plant species in the world.

 

Species that adapt the most aggressively to conditions in the Everglades, by spreading quickly or competing with native species that sometimes are threatened or endangered, are called "invasive". Thousands of exotic plant species have been observed in South Florida, usually introduced as ornamental landscaping, but park staff must eradicate such invasive plants as melaleuca tree (Melaleuca quinquenervia), Brazilian pepper (Schinus terebinthifolius), and Old World climbing fern (Lygodium microphyllum). Similarly, animals often do not find the predators or natural barriers to reproduction in the Everglades as they do where they originate, thus they often reproduce more quickly and efficiently. Lobate lac scale insects (Paratachardina pseudolobata) kill shrubs and other plants in hardwood hammocks. Bromeliad beetles (Metamasius callizona) destroy bromeliads and the ecosystems they host.

 

Walking catfish (Clarias batrachus) can deplete aquaculture stocks and they carry enteric septicemia. The Florida Fish and Wildlife Conservation Commission (FWC) listed eight "Reptiles of Concern", including the Burmese python (Python molurus bivittatus), focusing on them for their large sizes and aggressive natures, allowing licensed hunters to kill any listed animals in protected areas and sell their meat and hides. Burmese pythons, two subspecies of African rock pythons (Python sebae; northern and southern), and yellow anacondas (Eunectes notaeus) were banned from import into the U.S. in 2012. United States Secretary of the Interior Ken Salazar announced the inclusion of these reptiles at Everglades National Park. Exotic species control falls under the management of the U.S. Fish and Wildlife Service, which has been compiling and disseminating information about invasive species since 1994. Control of invasive species costs $500 million per year, but 1,700,000 acres (2,656.2 sq mi; 6,879.7 km2) of land in South Florida remains infested.

Elephants may be famed for their wrinkly exterior, but when it comes to Gillian Higgins' sculpture, it really is what’s on the inside that counts. This design, created by anatomist Gillian, depicts the intricate, interdependent and wonder of the internal systems on which life depends. Gillian believes that having a deeper understanding of our inner workings is essential in helping us to make wise decisions about our health and improve personal well being.

 

Designed by: Gillian Higgins

Gillian Higgins is the director of Horses Inside Out, an organisation which shows how understanding anatomy, physiology and bio mechanics can improve well being, performance and reduce risk of injury. Gillian’s unique trademark is painting the anatomical systems on the sides of live horses which she then uses as live canvases. Initially trained as a remedial sports and massage therapist, Gillian is also a McTimoney animal practitioner, author, British Horse Society Senior Coach and a competitive rider. She is regularly invited to give talks and demonstrations all over the world and has visited Australia, South Africa, North and South America as well as Europe and the British Isles.

 

Sponsored by: B Braun

Auction Price: £6000

 

Summer 2016, a herd of elephant sculptures descended on Sheffield for the biggest public art event the city has ever seen!

58 elephant sculptures, each uniquely decorated by artists, descended on Sheffield’s parks and open spaces, creating one of the biggest mass participation arts events the city has ever seen. Did you find them all?

The trail of elephants celebrates Sheffield’s creativity with over 75% of artists from the city. Some well-known names include Pete McKee, James Green, Jonathan Wilkinson and Lydia Monks – each of which has put their own creative mark on a 1.6m tall fibreglass elephant sculpture. They are all very difference, take a selfie with your favourite as they will be on display until the end of September.

International artist Mark Alexander, who is currently working with Rembrandt for an exhibition in Berlin, flew to Sheffield especially to paint his elephant and international players from the World Snooker Championship signed SnookHerd, an elephant celebrating the heritage of snooker in Sheffield.

The Arctic Monkeys, famous for their love of their home city, added their signatures to their own personalised sculpture which pays homage to the striking sound wave cover of the band’s 2013 album “AM”.

By supporting the Herd of Sheffield you are investing in the future of Sheffield Children’s Hospital. Every penny raised will go towards our Artfelt programme, which transforms the hospital’s walls and spaces with bright art, helping children recover in an environment tailored to them. The programme also puts on workshops for youngsters to provide distraction during anxious moments – such as before an operation, and to breakup long stays on the wards.

This exciting Wild in Art event brought to you by The Children’s Hospital Charity will:

Unite our city – bringing businesses, communities, artists, individuals and schools together to create a FREE sculpture trail which is accessible to all.

Attract more visitors – both nationally and regionally as well as encouraging thousands of people to become a tourist in their own city.

Invest in the future – with a city wide education programme that can be used for years to come and by funding a life-saving piece of medical equipment at Sheffield Children’s Hospital from the Herd auction at the end of the trail.

Showcase our city – celebrating Sheffield’s heritage and cementing our status as a vibrant and culturally exciting city through this world-class initiative.

 

The Herd of Sheffield Farewell Weekend was held on 14-16 October and was your chance to say a last goodbye to all 58 large elephant sculptures as they gather in one place for a final send-off at Meadowhall.

This special event gave visitors a chance to see the entire herd in all its glory – from the signed Arctic Monkeys’ ‘AM’ elephant, right through to ‘SnookHerd’, autographed by a host of international snooker players including current world champion Mark Selby.

Please note that the Little Herd elephants will not be on display as they will be returned to their school for pupils to enjoy.

Meadowhall, along with its joint owners, British Land are very proud to be supporting The Children’s Hospital Charity as host sponsors for the Herd of Sheffield Farewell Weekend.

 

Auction: Hundreds of elephant enthusiasts gathered at the Crucible on 20 October for the Herd of Sheffield Auction, which raised a total of £410,600 for The Children’s Hospital Charity.

HINDU GODDESS KALI

Kālī, also known as Kālikā (Sanskrit: कालिका), is the Hindu goddess associated with empowerment, shakti. She is the fierce aspect of the goddess Durga (Parvati). The name Kali comes from kāla, which means black, time, death, lord of death: Shiva. Since Shiva is called Kāla— the eternal time — the name of Kālī, his consort, also means "Time" or "Death" (as in "time has come"). Hence, Kāli is the Goddess of Time and Change. Although sometimes presented as dark and violent, her earliest incarnation as a figure of annihilation of evil forces still has some influence. Various Shakta Hindu cosmologies, as well as Shākta Tantric beliefs, worship her as the ultimate reality or Brahman. Comparatively recent devotional movements largely conceive Kāli as a benevolent mother goddess. Kālī is represented as the consort of Lord Shiva, on whose body she is often seen standing. Shiva lies in the path of Kali, whose foot on Shiva subdues her anger.

 

ETYMOLOGY

Kālī is the feminine form of kālam ("black, dark coloured"). Kāla primarily means "time" but also means "black" in honor of being the first creation before light itself. Kālī means "the black one" and refers to her being the entity of "time" or "beyond time." Kāli is strongly associated with Shiva, and Shaivas derive the masculine Kāla (an epithet of Shiva) to come from her feminine name. A nineteenth-century Sanskrit dictionary, the Shabdakalpadrum, states: कालः शिवः। तस्य पत्नीति - काली। kālaḥ śivaḥ। tasya patnīti kālī - "Shiva is Kāla, thus, his consort is Kāli" referring to Devi Parvathi being a manifestation of Devi MahaKali.

 

Other names include Kālarātri ("black night"), as described above, and Kālikā ("relating to time"). Coburn notes that the name Kālī can be used as a proper name, or as a description of color.

 

Kāli's association with darkness stands in contrast to her consort, Shiva, who manifested after her in creation, and who symbolises the rest of creation after Time is created. In his supreme awareness of Maya, his body is covered by the white ashes of the cremation ground (Sanskrit: śmaśāna) where he meditates, and with which Kāli is also associated, as śmaśāna-kālī.

 

ORIGINS

Hugh Urban notes that although the word Kālī appears as early as the Atharva Veda, the first use of it as a proper name is in the Kathaka Grhya Sutra (19.7). Kali is the name of one of the seven tongues of Agni, the [Rigvedic] God of Fire, in the Mundaka Upanishad (2:4), but it is unlikely that this refers to the goddess. The first appearance of Kāli in her present form is in the Sauptika Parvan of the Mahabharata (10.8.64). She is called Kālarātri (literally, "black night") and appears to the Pandava soldiers in dreams, until finally she appears amidst the fighting during an attack by Drona's son Ashwatthama. She most famously appears in the sixth century Devi Mahatmyam as one of the shaktis of Mahadevi, and defeats the demon Raktabija ("Bloodseed"). The tenth-century Kalika Purana venerates Kāli as the ultimate reality.

 

According to David Kinsley, Kāli is first mentioned in Hinduism as a distinct goddess around 600 CE, and these texts "usually place her on the periphery of Hindu society or on the battlefield." She is often regarded as the Shakti of Shiva, and is closely associated with him in various Puranas. The Kalika Purana depicts her as the "Adi Shakti" (Fundamental Power) and "Para Prakriti" or beyond nature.

 

WORSHIP & MANTRA

Kali could be considered a general concept, like Durga, and is mostly worshiped in the Kali Kula sect of worship. The closest way of direct worship is Maha Kali or Bhadra Kali (Bhadra in Sanskrit means 'gentle'). Kali is worshiped as one of the 10 Mahavidya forms of Adi Parashakti (Goddess Durga) or Bhagavathy according to the region. The mantra for worship is called Devi Argala Stotram.

Sanskrit: सर्वमङ्गलमाङ्गल्ये शिवे सर्वार्थसाधिके । शरण्ये त्र्यम्बके गौरि नारायणि नमोऽस्तु ते ॥

 

ॐ जयंती मंगल काली भद्रकाली कपालिनी । दुर्गा क्षमा शिवा धात्री स्वाहा स्वधा नमोऽस्तु‍ते ॥

(Sarvamaṅgalamāṅgalyē śivē sarvārthasādhikē . śaraṇyē tryambakē gauri nārāyaṇi namō'stu tē.

Oṃ jayantī mangala kālī bhadrakālī kapālinī . durgā kṣamā śivā dhātrī svāhā svadhā namō'stu‍tē.)

 

TANTRA

Goddesses play an important role in the study and practice of Tantra Yoga, and are affirmed to be as central to discerning the nature of reality as are the male deities. Although Parvati is often said to be the recipient and student of Shiva's wisdom in the form of Tantras, it is Kāli who seems to dominate much of the Tantric iconography, texts, and rituals. In many sources Kāli is praised as the highest reality or greatest of all deities. The Nirvana-tantra says the gods Brahma, Vishnu, and Shiva all arise from her like bubbles in the sea, ceaselessly arising and passing away, leaving their original source unchanged. The Niruttara-tantra and the Picchila-tantra declare all of Kāli's mantras to be the greatest and the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra all proclaim Kāli vidyas (manifestations of Mahadevi, or "divinity itself"). They declare her to be an essence of her own form (svarupa) of the Mahadevi.

 

In the Mahanirvana-tantra, Kāli is one of the epithets for the primordial sakti, and in one passage Shiva praises her:

 

At the dissolution of things, it is Kāla [Time] Who will devour all, and by reason of this He is called Mahākāla [an epithet of Lord Shiva], and since Thou devourest Mahākāla Himself, it is Thou who art the Supreme Primordial Kālika. Because Thou devourest Kāla, Thou art Kāli, the original form of all things, and because Thou art the Origin of and devourest all things Thou art called the Adya [the Primordial One]. Re-assuming after Dissolution Thine own form, dark and formless, Thou alone remainest as One ineffable and inconceivable. Though having a form, yet art Thou formless; though Thyself without beginning, multiform by the power of Maya, Thou art the Beginning of all, Creatrix, Protectress, and Destructress that Thou art.

 

The figure of Kāli conveys death, destruction, and the consuming aspects of reality. As such, she is also a "forbidden thing", or even death itself. In the Pancatattva ritual, the sadhaka boldly seeks to confront Kali, and thereby assimilates and transforms her into a vehicle of salvation. This is clear in the work of the Karpuradi-stotra, a short praise of Kāli describing the Pancatattva ritual unto her, performed on cremation grounds. (Samahana-sadhana)

 

He, O Mahākāli who in the cremation-ground, naked, and with dishevelled hair, intently meditates upon Thee and recites Thy mantra, and with each recitation makes offering to Thee of a thousand Akanda flowers with seed, becomes without any effort a Lord of the earth. Oh Kāli, whoever on Tuesday at midnight, having uttered Thy mantra, makes offering even but once with devotion to Thee of a hair of his Shakti [his energy/female companion] in the cremation-ground, becomes a great poet, a Lord of the earth, and ever goes mounted upon an elephant.

 

The Karpuradi-stotra clearly indicates that Kāli is more than a terrible, vicious, slayer of demons who serves Durga or Shiva. Here, she is identified as the supreme mistress of the universe, associated with the five elements. In union with Lord Shiva, she creates and destroys worlds. Her appearance also takes a different turn, befitting her role as ruler of the world and object of meditation. In contrast to her terrible aspects, she takes on hints of a more benign dimension. She is described as young and beautiful, has a gentle smile, and makes gestures with her two right hands to dispel any fear and offer boons. The more positive features exposed offer the distillation of divine wrath into a goddess of salvation, who rids the sadhaka of fear. Here, Kali appears as a symbol of triumph over death.

 

BENGALI TRADITION

Kali is also a central figure in late medieval Bengali devotional literature, with such devotees as Ramprasad Sen (1718–75). With the exception of being associated with Parvati as Shiva's consort, Kāli is rarely pictured in Hindu legends and iconography as a motherly figure until Bengali devotions beginning in the early eighteenth century. Even in Bengāli tradition her appearance and habits change little, if at all.

 

The Tantric approach to Kāli is to display courage by confronting her on cremation grounds in the dead of night, despite her terrible appearance. In contrast, the Bengali devotee appropriates Kāli's teachings adopting the attitude of a child, coming to love her unreservedly. In both cases, the goal of the devotee is to become reconciled with death and to learn acceptance of the way that things are. These themes are well addressed in Rāmprasād's work. Rāmprasād comments in many of his other songs that Kāli is indifferent to his wellbeing, causes him to suffer, brings his worldly desires to nothing and his worldly goods to ruin.

 

To be a child of Kāli, Rāmprasād asserts, is to be denied of earthly delights and pleasures. Kāli is said to refrain from giving that which is expected. To the devotee, it is perhaps her very refusal to do so that enables her devotees to reflect on dimensions of themselves and of reality that go beyond the material world.

 

A significant portion of Bengali devotional music features Kāli as its central theme and is known as Shyama Sangeet ("Music of the Night"). Mostly sung by male vocalists, today even women have taken to this form of music. One of the finest singers of Shyāma Sāngeet is Pannalal Bhattacharya.

 

In Bengal, Kāli is venerated in the festival Kali Puja, the new moon day of Ashwin month which coincides with Diwali festival.

 

In a unique form of Kāli worship, Shantipur worships Kāli in the form of a hand painted image of the deity known as Poteshwari (meaning the deity drawn on a piece of cloth).

 

LEGENDS

SLAYER OF RAKTABIJA

In Kāli's most famous legend, Devi Durga (Adi Parashakti) and her assistants, the Matrikas, wound the demon Raktabija, in various ways and with a variety of weapons in an attempt to destroy him. They soon find that they have worsened the situation for with every drop of blood that is dripped from Raktabija he reproduces a clone of himself. The battlefield becomes increasingly filled with his duplicates. Durga, in need of help, summons Kāli to combat the demons. It is said, in some versions, that Goddess Durga actually assumes the form of Goddess Kāli at this time. The Devi Mahatmyam describes:

 

Out of the surface of her (Durga's) forehead, fierce with frown, issued suddenly Kali of terrible countenance, armed with a sword and noose. Bearing the strange khatvanga (skull-topped staff ), decorated with a garland of skulls, clad in a tiger's skin, very appalling owing to her emaciated flesh, with gaping mouth, fearful with her tongue lolling out, having deep reddish eyes, filling the regions of the sky with her roars, falling upon impetuously and slaughtering the great asuras in that army, she devoured those hordes of the foes of the devas.

 

Kali destroys Raktabija by sucking the blood from his body and putting the many Raktabija duplicates in her gaping mouth. Pleased with her victory, Kali then dances on the field of battle, stepping on the corpses of the slain. In the Devi Mahatmya version of this story, Kali is also described as a Matrika and as a Shakti or power of Devi. She is given the epithet Cāṃuṇḍā (Chamunda), i.e. the slayer of the demons Chanda and Munda. Chamunda is very often identified with Kali and is very much like her in appearance and habit.

 

DAKSHINA KALI

In her most famous pose as Daksinakali, popular legends say that Kali, becoming drunk on the blood of her victims on the battlefield, dances with destructive frenzy. She is about to destroy the whole universe when, urged by all the gods, Shiva lies in her way to stop her. In her fury, she fails to see the body of Shiva lying amongst the corpses on the battlefield and steps upon his chest. Realizing Shiva lies beneath her feet, her anger is pacified and she calms her fury. Though not included in any of the puranas, popular legends state that Kali was ashamed at the prospect of keeping her husband beneath her feet and thus stuck her tongue out in shame. The Devi-Bhagavata Purana, which goes into great depths about the goddess Kali, reveals the tongue's actual symbolism.

 

The characteristic icons that depict Kali are the following; unbridled matted hair, open blood shot eyes, open mouth and a drooping tongue; in her hands, she holds a Khadga (bent sword or scimitar) and a human head; she has a girdle of human hands across her waist and an enchanted Shiva lies beneath her feet. Each of these icons represent a deep philosophical epithet. The drooping out-stuck tongue represents her blood-thirst. Lord Shiva beneath her feet represents matter, as Kali is undoubtedly the primeval energy. The depiction of Kali on Shiva shows that without energy, matter lies "dead". This concept has been simplified to a folk-tale depicting a wife placing her foot on her husband and sticking her tongue out in shame. In tantric contexts, the tongue is seen to denote the element (guna) of rajas (energy and action) controlled by sattva.

 

If Kali steps on Shiva with her right foot and holds the sword in her left hand, she is considered to be Dakshina Kali. The Dakshina Kali Temple has important religious associations with the Jagannath Temple and it is believed that Daksinakali is the guardian of the kitchen of the Lord Jagannath Temple. Puranic tradition says that in Puri, Lord Jagannath is regarded as Daksinakalika. Goddess Dakshinakali plays an important role in the 'Niti' of Saptapuri Amavasya.

 

One South Indian tradition tells of a dance contest between Shiva and Kali. After defeating the two demons Sumbha and Nisumbha, Kali takes up residence in the forest of Thiruvalankadu or Thiruvalangadu. She terrorizes the surrounding area with her fierce, disruptive nature. One of Shiva's devotees becomes distracted while performing austerities, and asks Shiva to rid the forest of the destructive goddess. When Shiva arrives, Kali threatens him, claiming the territory as her own. Shiva challenges Kali to a dance contest; both of them dance and Kali matches Shiva in every step that he takes until Shiva takes the "Urdhvatandava" step, by vertically raising his right leg. Kali refuses to perform this step, which would not befit her as a woman, and became pacified.

 

SMASHAN KALI

If the Kali steps out with the left foot and holds the sword in her right hand, she is the terrible form of Mother, the Smashan Kali of the cremation ground. She is worshiped by tantrics, the followers of Tantra, who believe that one's spiritual discipline practiced in a smashan (cremation ground) brings success quickly. Sarda Devi, the consort of Ramakrishna Paramhansa, worshipped Smashan Kali at Dakshineshwar.

 

MATERNAL KALI

Another legend depicts the infant Shiva calming Kali. In this similar story, Kali has defeated her enemies on the battlefield and begun to dance out of control, drunk on the blood of the slain. To calm her down and to protect the stability of the world, Shiva is sent to the battlefield, as an infant, crying aloud. Seeing the child's distress, Kali ceases dancing to care for the helpless infant. She picks him up, kisses his head, and proceeds to breast feed the infant Shiva. This legend is notable because it shows Kali in her benevolent, maternal aspect, with which she is not usually identified.

 

MAHAKALI

Mahakali (Sanskrit: Mahākālī, Devanagari: महाकाली), literally translated as Great Kali, is sometimes considered as a greater form of Kali, identified with the Ultimate reality of Brahman. It can also be used as an honorific of the Goddess Kali, signifying her greatness by the prefix "Mahā-". Mahakali, in Sanskrit, is etymologically the feminized variant of Mahakala or Great Time (which is interpreted also as Death), an epithet of the God Shiva in Hinduism. Mahakali is the presiding Goddess of the first episode of the Devi Mahatmya. Here she is depicted as Devi in her universal form as Shakti. Here Devi serves as the agent who allows the cosmic order to be restored.

 

Kali is depicted in the Mahakali form as having ten heads, ten arms, and ten legs. Each of her ten hands is carrying a various implement which vary in different accounts, but each of these represent the power of one of the Devas or Hindu Gods and are often the identifying weapon or ritual item of a given Deva. The implication is that Mahakali subsumes and is responsible for the powers that these deities possess and this is in line with the interpretation that Mahakali is identical with Brahman. While not displaying ten heads, an "ekamukhi" or one headed image may be displayed with ten arms, signifying the same concept: the powers of the various Gods come only through Her grace.

 

ICONOGRAPHY

Kali is portrayed mostly in two forms: the popular four-armed form and the ten-armed Mahakali form. In both of her forms, she is described as being black in color but is most often depicted as blue in popular Indian art. Her eyes are described as red with intoxication, and in absolute rage, her hair is shown disheveled, small fangs sometimes protrude out of her mouth, and her tongue is lolling. She is often shown naked or just wearing a skirt made of human arms and a garland of human heads. She is also accompanied by serpents and a jackal while standing on a seemingly dead Shiva, usually right foot forward to symbolize the more popular Dakshinamarga or right-handed path, as opposed to the more infamous and transgressive Vamamarga or left-handed path.

 

In the ten-armed form of Mahakali she is depicted as shining like a blue stone. She has ten faces and ten feet and three eyes. She has ornaments decked on all her limbs. There is no association with Shiva.

 

The Kalika Purana describes Kali as possessing a soothing dark complexion, as perfectly beautiful, riding a lion, four-armed, holding a sword and blue lotuses, her hair unrestrained, body firm and youthful.

 

In spite of her seemingly terrible form, Kali Ma is often considered the kindest and most loving of all the Hindu goddesses, as she is regarded by her devotees as the Mother of the whole Universe. And because of her terrible form, she is also often seen as a great protector. When the Bengali saint Ramakrishna once asked a devotee why one would prefer to worship Mother over him, this devotee rhetorically replied, "Maharaj, when they are in trouble your devotees come running to you. But, where do you run when you are in trouble?"

 

According to Ramakrishna, darkness is the Ultimate Mother, or Kali:

 

My Mother is the principle of consciousness. She is Akhanda Satchidananda; indivisible Reality, Awareness, and Bliss. The night sky between the stars is perfectly black. The waters of the ocean depths are the same; The infinite is always mysteriously dark. This inebriating darkness is my beloved Kali.

 

SRI RAMAKRISHNA

This is clear in the works of such contemporary artists as Charles Wish, and Tyeb Mehta, who sometimes take great liberties with the traditional, accepted symbolism, but still demonstrate a true reverence for the Shakta sect.

 

POPULAR FORM

Classic depictions of Kali share several features, as follows:

 

Kali's most common four armed iconographic image shows each hand carrying variously a sword, a trishul (trident), a severed head and a bowl or skull-cup (kapala) catching the blood of the severed head.

 

Two of these hands (usually the left) are holding a sword and a severed head. The Sword signifies Divine Knowledge and the Human Head signifies human Ego which must be slain by Divine Knowledge in order to attain Moksha. The other two hands (usually the right) are in the abhaya (fearlessness) and varada (blessing) mudras, which means her initiated devotees (or anyone worshipping her with a true heart) will be saved as she will guide them here and in the hereafter.

 

She has a garland consisting of human heads, variously enumerated at 108 (an auspicious number in Hinduism and the number of countable beads on a Japa Mala or rosary for repetition of Mantras) or 51, which represents Varnamala or the Garland of letters of the Sanskrit alphabet, Devanagari. Hindus believe Sanskrit is a language of dynamism, and each of these letters represents a form of energy, or a form of Kali. Therefore she is generally seen as the mother of language, and all mantras.

 

She is often depicted naked which symbolizes her being beyond the covering of Maya since she is pure (nirguna) being-consciousness-bliss and far above prakriti. She is shown as very dark as she is brahman in its supreme unmanifest state. She has no permanent qualities - she will continue to exist even when the universe ends. It is therefore believed that the concepts of color, light, good, bad do not apply to her - she is the pure, un-manifested energy, the Adi-shakti.

 

SHIVA IN KALI ICONOGRAPHY

In both these images she is shown standing on the prone, inert or dead body of Shiva. There is a legend for the reason behind her standing on what appears to be Shiva's corpse, which translates as follows:

 

Once Kali had destroyed all the demons in battle, she began a terrific dance out of the sheer joy of victory. All the worlds or lokas began to tremble and sway under the impact of her dance. So, at the request of all the Gods, Shiva himself asked her to desist from this behavior. However, she was too intoxicated to listen. Hence, Shiva lay like a corpse among the slain demons in order to absorb the shock of the dance into himself. When Kali eventually stepped upon Shiva, she realized she was trampling and hurting her husband and bit her tongue in shame.

 

The story described here is a popular folk tale and not described or hinted in any of the puranas. The puranic interpretation is as follows:

 

Once, Parvati asks Shiva to chose the one form among her 10 forms which he likes most. To her surprise, Shiva reveals that he is most comfortable with her Kali form, in which she is bereft of her jewellery, her human-form, her clothes, her emotions and where she is only raw, chaotic energy, where she is as terrible as time itself and even greater than time. As Parvati takes the form of Kali, Shiva lies at her feet and requests her to place her foot on his chest, upon his heart. Once in this form, Shiva requests her to have this place, below her feet in her iconic image which would be worshiped throughout.

 

This idea has been explored in the Devi-Bhagavata Purana and is most popular in the Shyama Sangeet, devotional songs to Kali from the 12th to 15th centuries.

 

The Tantric interpretation of Kali standing on top of her husband is as follows:

 

The Shiv tattava (Divine Consciousness as Shiva) is inactive, while the Shakti tattava (Divine Energy as Kali) is active. Shiva and Kali represent Brahman, the Absolute pure consciousness which is beyond all names, forms and activities. Kali, on the other hand, represents the potential (and manifested) energy responsible for all names, forms and activities. She is his Shakti, or creative power, and is seen as the substance behind the entire content of all consciousness. She can never exist apart from Shiva or act independently of him, just as Shiva remains a mere corpse without Kali i.e., Shakti, all the matter/energy of the universe, is not distinct from Shiva, or Brahman, but is rather the dynamic power of Brahman. Hence, Kali is Para Brahman in the feminine and dynamic aspect while Shiva is the male aspect and static. She stands as the absolute basis for all life, energy and beneath her feet lies, Shiva, a metaphor for mass, which cannot retain its form without energy.

 

While this is an advanced concept in monistic Shaktism, it also agrees with the Nondual Trika philosophy of Kashmir, popularly known as Kashmir Shaivism and associated most famously with Abhinavagupta. There is a colloquial saying that "Shiva without Shakti is Shava" which means that without the power of action (Shakti) that is Mahakali (represented as the short "i" in Devanagari) Shiva (or consciousness itself) is inactive; Shava means corpse in Sanskrit and the play on words is that all Sanskrit consonants are assumed to be followed by a short letter "a" unless otherwise noted. The short letter "i" represents the female power or Shakti that activates Creation. This is often the explanation for why She is standing on Shiva, who is either Her husband and complement in Shaktism or the Supreme Godhead in Shaivism.

 

To properly understand this complex Tantric symbolism it is important to remember that the meaning behind Shiva and Kali does not stray from the non-dualistic parlance of Shankara or the Upanisads. According to both the Mahanirvana and Kularnava Tantras, there are two distinct ways of perceiving the same absolute reality. The first is a transcendental plane which is often described as static, yet infinite. It is here that there is no matter, there is no universe and only consciousness exists. This form of reality is known as Shiva, the absolute Sat-Chit-Ananda - existence, knowledge and bliss. The second is an active plane, an immanent plane, the plane of matter, of Maya, i.e., where the illusion of space-time and the appearance of an actual universe does exist. This form of reality is known as Kali or Shakti, and (in its entirety) is still specified as the same Absolute Sat-Chit-Ananda. It is here in this second plane that the universe (as we commonly know it) is experienced and is described by the Tantric seer as the play of Shakti, or God as Mother Kali.

 

From a Tantric perspective, when one meditates on reality at rest, as absolute pure consciousness (without the activities of creation, preservation or dissolution) one refers to this as Shiva or Brahman. When one meditates on reality as dynamic and creative, as the Absolute content of pure consciousness (with all the activities of creation, preservation or dissolution) one refers to it as Kali or Shakti. However, in either case the yogini or yogi is interested in one and the same reality - the only difference being in name and fluctuating aspects of appearance. It is this which is generally accepted as the meaning of Kali standing on the chest of Shiva.

 

Although there is often controversy surrounding the images of divine copulation, the general consensus is benign and free from any carnal impurities in its substance. In Tantra the human body is a symbol for the microcosm of the universe; therefore sexual process is responsible for the creation of the world. Although theoretically Shiva and Kali (or Shakti) are inseparable, like fire and its power to burn, in the case of creation they are often seen as having separate roles. With Shiva as male and Kali as female it is only by their union that creation may transpire. This reminds us of the prakrti and purusa doctrine of Samkhya wherein prakāśa- vimarśa has no practical value, just as without prakrti, purusa is quite inactive. This (once again) stresses the interdependencies of Shiva and Shakti and the vitality of their union.

 

Gopi Krishna proposed that Kali standing on the dead Shiva or Shava (Sanskrit for dead body) symbolised the helplessness of a person undergoing the changing process (psychologically and physiologically) in the body conducted by the Kundalini Shakti.

 

DEVELOPMENT

In the later traditions, Kali has become inextricably linked with Shiva. The unleashed form of Kali often becomes wild and uncontrollable, and only Shiva is able to tame her just as only Kali can tame Shiva. This is both because she is often a transformed version of one of his consorts and because he is able to match her wildness.

 

The ancient text of Kali Kautuvam describes her competition with Shiva in dance, from which the sacred 108 Karanas appeared. Shiva won the competition by acting the urdva tandava, one of the Karanas, by raising his feet to his head. Other texts describe Shiva appearing as a crying infant and appealing to her maternal instincts. While Shiva is said to be able to tame her, the iconography often presents her dancing on his fallen body, and there are accounts of the two of them dancing together, and driving each other to such wildness that the world comes close to unravelling.

 

Shiva's involvement with Tantra and Kali's dark nature have led to her becoming an important Tantric figure. To the Tantric worshippers, it was essential to face her Curse, the terror of death, as willingly as they accepted Blessings from her beautiful, nurturing, maternal aspect. For them, wisdom meant learning that no coin has only one side: as death cannot exist without life, so life cannot exist without death. Kali's role sometimes grew beyond that of a chaos - which could be confronted - to that of one who could bring wisdom, and she is given great metaphysical significance by some Tantric texts. The Nirvāna-tantra clearly presents her uncontrolled nature as the Ultimate Reality, claiming that the trimurti of Brahma, Vishnu and Rudra arise and disappear from her like bubbles from the sea. Although this is an extreme case, the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra declare her the svarupa (own-being) of the Mahadevi (the great Goddess, who is in this case seen as the combination of all devis).

 

The final stage of development is the worshipping of Kali as the Great Mother, devoid of her usual violence. This practice is a break from the more traditional depictions. The pioneers of this tradition are the 18th century Shakta poets such as Ramprasad Sen, who show an awareness of Kali's ambivalent nature. Ramakrishna, the 19th century Bengali saint, was also a great devotee of Kali; the western popularity of whom may have contributed to the more modern, equivocal interpretations of this Goddess. Rachel McDermott's work, however, suggests that for the common, modern worshipper, Kali is not seen as fearful, and only those educated in old traditions see her as having a wrathful component. Some credit to the development of Devi must also be given to Samkhya. Commonly referred to as the Devi of delusion, Mahamaya or Durga, acting in the confines of (but not being bound by) the nature of the three gunas, takes three forms: Maha-Kali, Maha-Lakshmi and Maha-Saraswati, being her tamas-ika, rajas-ika and sattva-ika forms. In this sense, Kali is simply part of a larger whole.

 

Like Sir John Woodroffe and Georg Feuerstein, many Tantric scholars (as well as sincere practitioners) agree that, no matter how propitious or appalling you describe them, Shiva and Devi are simply recognizable symbols for everyday, abstract (yet tangible) concepts such as perception, knowledge, space-time, causation and the process of liberating oneself from the confines of such things. Shiva, symbolizing pure, absolute consciousness, and Devi, symbolizing the entire content of that consciousness, are ultimately one and the same - totality incarnate, a micro-macro-cosmic amalgamation of all subjects, all objects and all phenomenal relations between the "two." Like man and woman who both share many common, human traits yet at the same time they are still different and, therefore, may also be seen as complementary.

 

Worshippers prescribe various benign and horrific qualities to Devi simply out of practicality. They do this so they may have a variety of symbols to choose from, symbols which they can identify and relate with from the perspective of their own, ever-changing time, place and personal level of unfolding. Just like modern chemists or physicists use a variety of molecular and atomic models to describe what is unperceivable through rudimentary, sensory input, the scientists of ontology and epistemology must do the same. One of the underlying distinctions of Tantra, in comparison to other religions, is that it allows the devotee the liberty to choose from a vast array of complementary symbols and rhetoric which suit one's evolving needs and tastes. From an aesthetic standpoint, nothing is interdict and nothing is orthodox. In this sense, the projection of some of Devi's more gentle qualities onto Kali is not sacrilege and the development of Kali really lies in the practitioner, not the murthi.

______________________

 

HINDU GODDESS DURGA

Durga, meaning "the inaccessible" or "the invincible", is the most popular incarnation of Devi and one of the main forms of the Goddess Shakti in the Hindu pantheon. Durga is the original manifested form of Mother Parvati or Adi-Parashakti. Durga is Adi-Parashakti herself. The Devi Gita, declares her to be the greatest Goddess. Thus, she is considered the supreme goddess and primary deity in Shaktism, occupying a place similar to Lord Krishna in Vaishnavism. According to Skanda Purana, the goddess Parvati accounted the name "Durga" after she killed the demon Durgamaasura. Goddess Parvati is considered to be the complete incarnation of Adi Parashakti or Goddess Durga, with all other goddesses being her incarnations or manifestations. Adi Parashakti or Mahadevi, the supreme power, is called Durga Shakti as per Devi-Mahatmya. Adi Parashakti or Devi Durga is a Hindu concept of the Ultimate Shakti or Mahashakti, the ultimate power inherent in all Creation. This is especially prevalent in the Shakta denomination within Hinduism, which worships the Goddess Devi in all her manifestations. She is Goddess Lakshmi and Goddess Saraswati in her mild form; Goddess Kali and Goddess Chandi in her wrathful form. Durga is also called Padmanabha-Sahodari and Narayani, the sister of Lord Vishnu. According to Shaivism and Shaktism She is supreme, but to bring back lord Shiva in Sansar, she was reborn in human form (Sati and Parvati) to marry Shiva. Durga gave birth to his first child Kartikeya.

 

ORIGINS & DEVELOPMENT

Ramprasad Chanda writes the following about the development of Durga from primitive goddess to her current form:

 

"...it is possible to distinguish two different strata – one primitive and the other advanced. The primitive form of Durga is the result of syncretism of a mountain-goddess worshiped by the dwellers of the Himalaya and the Vindhyas, a goddess worshiped by the nomadic Abhira shepherd, the vegetation spirit conceived as a female, and a war-goddess. As her votaries advanced in civilization the primitive war-goddess was transformed into the personification of the all-destroying time (Kali), the vegetation spirit into the primordial energy (Adya Sakti) and the saviouress from “samsara” (cycle of rebirths) , and gradually brought into line with the Brahmanic mythology and philosophy."

 

It is not possible to date her The delusion of the supreme soul is otherwise called Shakti (power). From this power, generates all forms of knowledge of the world and it is accepted as vital cause of creation, existence and destruction. According to 'Shree Durga Shaptshati- Rahasyam', the original power is Mahalaxmi that created three pairs of Supreme Powers. They are Brahma, Vishnu and Shiva as male and Saraswati, Laxmi and Parvati as female, and they married respectively. Maha Saraswati is well known as Brahmani, Mahalaxmi as Vaishnavi and Mahakali as Maheswari. Durga Shakti is the original cause of all the present or past worldly occurrences. Durga Shakti is called as Adhyashakti, Paramatma Shakti or Ati Prakrutika Shakti. She is creating and controlling other two powers: Natural and General. Natural Power is called as Atma Shakti, Prakrutika Shakti, Pancha Mahabhuta Shakti etc. This Shakti creates and controls the General Energy. General Energies are called Jada Shakti or Tamashakti. By the blessings of Durga Shakti, the mother of the Universe, man is able to get his emancipation or salvation and indulge in enjoyments in performance of his daily activities. So Vyasadev, the eminent poet of "Devi Bhagwat", has aptly described "Rudrahinam Vishnuhinam na vadanti janastatha Shaktihinam Yathasarbe probodhanti Naradhamam". The powerless persons are despised as mean persons. So, by being devoted to the Supreme, we should be strong and powerful by her grace.

 

STORIES

Shiva Purana gives an account of the origin of Durga. At the beginning of time, Lord Shiva invoked Durga, the primordial energy from his left half to create. Together they created their eternal abode, Shivaloka, also known as Kashi. Thereafter, they created Vishnu and Brahma.

 

As per Shiva Purana and Devi Mahatmyah, Mahishasura, the son of demon Rambha, unleashed reign of terror on earth. When gods intervened, Mahishasura defeated gods and banished them from heaven. Vanquished gods went to Trideva- Brahma, Vishnu and Shiva. As they narrated their woeful tale, immense mass of light manifested from Lord Vishnu's mouth, which was joined by similar rays that emerged from the enraged faces of gods. This mass of light transformed into a woman. Then all the Gods gave their devine weapon to that supreme power. Adishakti re-manifested as Durga to slay Mahishasura. Armed with celestial weapons of all deities and decked with divine ornaments, Durga rode into the battle field and challenged demons for battle. Mahishasura's entire army, led by demons like Chikshur, Chamar, Asiloma, Vidalaksha, Durdhara, Durmukha, Mahahanu and many more attacked Durga at once. But Durga slew all of them with unparalleled cruelty. An enraged Mahishasura attacked Durga in guise of a buffalo. But Durga bound it with ropes. The buffalo morphed into a lion and lept on Durga, but she beheaded it with her sword. At this, Mahishasura began to fight in form of a swordsman. Durga pinned him down with a torrent of arrows. Mahishasura now assumed form of a giant elephant and tugged at Durga's lion. Durga lopped off its trunk with her sword and freed her lion. The elephent turned into a buffalo and charged at Durga. Sipping from her wine cup, Durga flung her trident and beheaded Mahishasura, finally killing him.

 

WIKIPEDIA

From 1833 until 1932, the present day Oakes Garden Theatre was the site of the once famous Clifton Hotels. Both hotels, at different times, were destroyed by fire. Named in honor of the late Sir Harry Oakes, a Parks Commissioner who purchased the site and then gave the land to The Niagara Parks Commission in exchange for another plot of land, Oakes Garden Theatre construction began in 1935, with the formal opening taking place on September 18, 1937.

   

Oakes Garden Theatre - Niagara Parks, Canada

 

Oakes Garden Theatre is a highly decorative Beaux Arts landscape, including classical and picturesque elements, with the central point a grass Greco-Roman amphitheatre surrounded by a limestone staircase, and features fountains, urns, statuary and sculptured bas-relief panels and medallions. It is historically significant, as “very few examples of landscape architectural design of this age or complexity exist in Ontario or Canada.” (P. Berketo, NPC Study, 2000)

 

Oakes Garden Theatre “is fan-shaped, with the stage so placed that the panorama of the Falls forms a natural backdrop, while at the rear there is a curved pergola connecting two open pavilions, one oriented on the axis of the Horseshoe Falls, the other on that of the American Falls.” (NPC Annual Report, 1944) Behind the pergola, is a beautiful and serene Japanese-style garden with a lily pond and wishing bridge, which is often used as a back-drop for wedding and other photographs.

 

Designed by Howard Dunington-Grubb, who has been called the father of Canadian landscape architecture, and assisted by J.V. Stensson and architect William Somerville, Oakes Garden Theatre displays specifically outlined spaces, utilizing architecture and sculpture, to provide remarkable views and vistas.

 

Formal gardens using carpet bedding - a Victorian practice of forming beds of low-growing plants of the same height in geometric patterns - are laid out and interspersed with flagstone walkways and expanses of grass. The unique design takes advantage of the various levels on the site.

 

Florence Wyle, Frances Loring and Elizabeth Wyn Wood, all significant Canadian sculptors from Toronto, were commissioned to design the fountains and bas-relief sculptures within the garden. Their works are neo-classical, which treasures beauty, and due to the subject matter of indigenous plants and animals, are considered uniquely Canadian.

 

Surrounded by an ornamental stone wall, this surprising garden provides visitors with a feeling of fantasy and escape.

 

Initially, the theatre was used for band and choir concerts. More recently, it has been used for corporate receptions, movie screenings and live broadcasts of internationally televised programs.

 

Following the construction of the present Rainbow Bridge, the Niagara Falls Bridge Commission partnered with The Niagara Parks Commission in 1941 to create Rainbow Gardens as a link between the bridge and Oakes Garden Theatre and as an entrance to Canada and Queen Victoria Park. It was designed by the same team who designed Oakes Garden Theatre. “These gardens are equally as formal but have details typical of the Art Deco and Moderne period.” (P. Berketo, NPC Study, 2000)

 

Oakes Garden Theatre (Beaux Arts) and Rainbow Gardens (Art Deco) are two separate yet inter-penetrating spaces with areas of each interdependent on the other. Both gardens showcase early twentieth century Canadian art, architecture and landscape architecture.

A gorgeous photo of the Bridal Veil Falls in Provo Canyon, Utah.

 

If there's one aspect of the Earth that the USGS knows well...it's our water resources. Of course, USGS is a leader in​ providing data and information for​ ​many aspects of ​earth science, but we also know that water is key to our quality life. Clean, plentiful, healthy, and affordable water for future generations starts with understanding the availability of our water. A changing climate affects the quality and quantity of water, and understanding that effect leads to a ​more ​secure water future. Understanding floods and their characteristics is a matter of public safety. Even knowing the interdependent relationship between water and energy can lead to a securer energy future. For all these things, the USGS provides vital research and data to help ​people and decision-makers better understand the planet we inhabit and rely upon daily.

 

Learn more about our water resources and data at water.usgs.gov.

 

Photo credit: Ann-Marie Mair (ringlitephotography on Flickr)

In Chinese philosophy, yin and yang (also, yin-yang or yin yang) describes how apparently opposite or contrary forces are actually complementary, interconnected, and interdependent in the natural world, and how they give rise to each other as they interrelate to one another. Many tangible dualities (such as light and dark, fire and water, expanding and contracting) are thought of as physical manifestations of the duality symbolized by yin and yang. This duality lies at the origins (Wikipedia)

ho·me·o·sta·sis

ˌhōmēəˈstāsəs/

noun

the tendency toward a relatively stable equilibrium between interdependent elements, especially as maintained by physiological processes.

Elephants may be famed for their wrinkly exterior, but when it comes to Gillian Higgins' sculpture, it really is what’s on the inside that counts. This design, created by anatomist Gillian, depicts the intricate, interdependent and wonder of the internal systems on which life depends. Gillian believes that having a deeper understanding of our inner workings is essential in helping us to make wise decisions about our health and improve personal well being.

 

Designed by: Gillian Higgins

Gillian Higgins is the director of Horses Inside Out, an organisation which shows how understanding anatomy, physiology and bio mechanics can improve well being, performance and reduce risk of injury. Gillian’s unique trademark is painting the anatomical systems on the sides of live horses which she then uses as live canvases. Initially trained as a remedial sports and massage therapist, Gillian is also a McTimoney animal practitioner, author, British Horse Society Senior Coach and a competitive rider. She is regularly invited to give talks and demonstrations all over the world and has visited Australia, South Africa, North and South America as well as Europe and the British Isles.

 

Sponsored by: B Braun

Auction Price: £6000

 

Summer 2016, a herd of elephant sculptures descended on Sheffield for the biggest public art event the city has ever seen!

58 elephant sculptures, each uniquely decorated by artists, descended on Sheffield’s parks and open spaces, creating one of the biggest mass participation arts events the city has ever seen. Did you find them all?

The trail of elephants celebrates Sheffield’s creativity with over 75% of artists from the city. Some well-known names include Pete McKee, James Green, Jonathan Wilkinson and Lydia Monks – each of which has put their own creative mark on a 1.6m tall fibreglass elephant sculpture. They are all very difference, take a selfie with your favourite as they will be on display until the end of September.

International artist Mark Alexander, who is currently working with Rembrandt for an exhibition in Berlin, flew to Sheffield especially to paint his elephant and international players from the World Snooker Championship signed SnookHerd, an elephant celebrating the heritage of snooker in Sheffield.

The Arctic Monkeys, famous for their love of their home city, added their signatures to their own personalised sculpture which pays homage to the striking sound wave cover of the band’s 2013 album “AM”.

By supporting the Herd of Sheffield you are investing in the future of Sheffield Children’s Hospital. Every penny raised will go towards our Artfelt programme, which transforms the hospital’s walls and spaces with bright art, helping children recover in an environment tailored to them. The programme also puts on workshops for youngsters to provide distraction during anxious moments – such as before an operation, and to breakup long stays on the wards.

This exciting Wild in Art event brought to you by The Children’s Hospital Charity will:

Unite our city – bringing businesses, communities, artists, individuals and schools together to create a FREE sculpture trail which is accessible to all.

Attract more visitors – both nationally and regionally as well as encouraging thousands of people to become a tourist in their own city.

Invest in the future – with a city wide education programme that can be used for years to come and by funding a life-saving piece of medical equipment at Sheffield Children’s Hospital from the Herd auction at the end of the trail.

Showcase our city – celebrating Sheffield’s heritage and cementing our status as a vibrant and culturally exciting city through this world-class initiative.

 

The Herd of Sheffield Farewell Weekend was held on 14-16 October and was your chance to say a last goodbye to all 58 large elephant sculptures as they gather in one place for a final send-off at Meadowhall.

This special event gave visitors a chance to see the entire herd in all its glory – from the signed Arctic Monkeys’ ‘AM’ elephant, right through to ‘SnookHerd’, autographed by a host of international snooker players including current world champion Mark Selby.

Please note that the Little Herd elephants will not be on display as they will be returned to their school for pupils to enjoy.

Meadowhall, along with its joint owners, British Land are very proud to be supporting The Children’s Hospital Charity as host sponsors for the Herd of Sheffield Farewell Weekend.

 

Auction: Hundreds of elephant enthusiasts gathered at the Crucible on 20 October for the Herd of Sheffield Auction, which raised a total of £410,600 for The Children’s Hospital Charity.

Hōʻawa or Kona cheesewood

Pittosporaceae (Pittosporum family)

Endemic to the Hawaiian Islands (Kohala Mountains south to the Kaʻū District, Hawaiʻi Island)

Oʻahu (Cultivated)

 

The Hawaiian crow, or ʻalalā, (Corvus tropicus) fed on this hōʻawa, attracted by the bright orange inside color of the ripe fruit capsules and dark seeds and thus ensuring the spread of the plants. However, ʻalalā are now extremely rare and populations of hōʻawa in the birds former range are becoming very scarce as a result. This is a classic example of how interdependent native Hawaiian plants and animals are in the natural ecosystem.

 

The early Hawaiians used the wood to make gunwales for canoes.

The outer layer of the fruit valves were used medicinally. They were pounded and used externally on sores.

 

Flowers

www.flickr.com/photos/dweickhoff/5187804829/in/photolist-...

 

Leaves

www.flickr.com/photos/dweickhoff/8054148306/in/photolist-...

 

Habit

www.flickr.com/photos/dweickhoff/5188406532/in/photolist-...

 

Etymology

The generic name Pittosporum is derived from the Greek pittos, pitch, and sporos, seed, in reference to the film of viscid resin covering the black seeds.

 

The specific epithet hosmeri is named in honor of Ralph Sheldon Hosmer (1874-1963), Hawaiʻi's first territorial forester. Hosmer's Grove, Maui is one of Hosmer's forestry experimental using non-native species he planted in 1910. Unfortunately, some of the species are now invasive on Maui, such as the Mexican weeping pine, Monterrey pine, and eucalyptus, displacing the native plants.

"Merce Cunningham: Common Time" is a major retrospective exhibition organized by the Walker Art Center that is appearing simultaneously at the Walker and the MCA. Merce Cunningham (American, 1919–2009) was a seminal figure in modern dance, revolutionizing performance through his choreography and world-renowned dance company and through partnerships with leading artists, who created costumes, lighting, and set designs for his company’s performances.

 

The exhibition showcases Cunningham’s multidisciplinary projects, exploring, as Cunningham described, the “underlying principle that music and dance and art could be separate entities independent and interdependent, sharing a common time.”

The Everglades are subtropical wetlands located in the southern portion of the U.S. state of Florida, comprising the southern half of a large watershed. The system begins near Orlando with the Kissimmee River, which discharges into the vast but shallow Lake Okeechobee. Water leaving the lake in the wet season forms a slow-moving river 60 miles (97 km) wide and over 100 miles (160 km) long, flowing southward across a limestone shelf to Florida Bay at the southern end of the state. The Everglades are shaped by water and fire, experiencing frequent flooding in the wet season and drought in the dry season. Writer Marjory Stoneman Douglas popularized the term “River of Grass” to describe the sawgrass marshes, part of a complex system of interdependent ecosystems that include cypress swamps, the estuarine mangrove forests of the Ten Thousand Islands, tropical hardwood hammocks, pine rockland, and the marine environment of Florida Bay. Wikipedia.

vdostalek.wordpress.com/2010/04/14/lonely-alligator/

Last night was cold. So cold I wanted to stay in, yet the light called me to the halo hanging high in the sky. I went out into the yard and it was beautiful, and the light called me further. Into the car and down the hill not far from from home. Fantastic clouds stretching across the sky. In the field, the tree always teasing me with it's ideas. I always thought of it as two halves of brain. A lot of ideas come from this place. The light calls further. I had never walked to the tree, but here I am in the dead of winter. The tree of ideas calls. Ever closer the tree brings me. I call it one tree because that is what it is to me, symbiotic, intertwined, interdependent, like life. This is the Good Life. I had the pleasure of sharing this image on facebook and see it speak to me in a similar manner and deeper. It has reminded me deeply of spiritual things. I thank all of you for sharing your words.

 

spirithands.smugmug.com/Landscapes/i-9fZRKcx/A

 

Kālī, also known as Kālikā (Sanskrit: कालिका), is the Hindu goddess associated with empowerment, shakti. She is the fierce aspect of the goddess Durga (Parvati). The name Kali comes from kāla, which means black, time, death, lord of death: Shiva. Since Shiva is called Kāla— the eternal time — the name of Kālī, his consort, also means "Time" or "Death" (as in "time has come"). Hence, Kāli is the Goddess of Time and Change. Although sometimes presented as dark and violent, her earliest incarnation as a figure of annihilation of evil forces still has some influence. Various Shakta Hindu cosmologies, as well as Shākta Tantric beliefs, worship her as the ultimate reality or Brahman. Comparatively recent devotional movements largely conceive Kāli as a benevolent mother goddess. Kālī is represented as the consort of Lord Shiva, on whose body she is often seen standing. Shiva lies in the path of Kali, whose foot on Shiva subdues her anger.

 

ETYMOLOGY

Kālī is the feminine form of kālam ("black, dark coloured"). Kāla primarily means "time" but also means "black" in honor of being the first creation before light itself. Kālī means "the black one" and refers to her being the entity of "time" or "beyond time." Kāli is strongly associated with Shiva, and Shaivas derive the masculine Kāla (an epithet of Shiva) to come from her feminine name. A nineteenth-century Sanskrit dictionary, the Shabdakalpadrum, states: कालः शिवः। तस्य पत्नीति - काली। kālaḥ śivaḥ। tasya patnīti kālī - "Shiva is Kāla, thus, his consort is Kāli" referring to Devi Parvathi being a manifestation of Devi MahaKali.

 

Other names include Kālarātri ("black night"), as described above, and Kālikā ("relating to time"). Coburn notes that the name Kālī can be used as a proper name, or as a description of color.

 

Kāli's association with darkness stands in contrast to her consort, Shiva, who manifested after her in creation, and who symbolises the rest of creation after Time is created. In his supreme awareness of Maya, his body is covered by the white ashes of the cremation ground (Sanskrit: śmaśāna) where he meditates, and with which Kāli is also associated, as śmaśāna-kālī.

 

ORIGINS

Hugh Urban notes that although the word Kālī appears as early as the Atharva Veda, the first use of it as a proper name is in the Kathaka Grhya Sutra (19.7). Kali is the name of one of the seven tongues of Agni, the [Rigvedic] God of Fire, in the Mundaka Upanishad (2:4), but it is unlikely that this refers to the goddess. The first appearance of Kāli in her present form is in the Sauptika Parvan of the Mahabharata (10.8.64). She is called Kālarātri (literally, "black night") and appears to the Pandava soldiers in dreams, until finally she appears amidst the fighting during an attack by Drona's son Ashwatthama. She most famously appears in the sixth century Devi Mahatmyam as one of the shaktis of Mahadevi, and defeats the demon Raktabija ("Bloodseed"). The tenth-century Kalika Purana venerates Kāli as the ultimate reality.

 

According to David Kinsley, Kāli is first mentioned in Hinduism as a distinct goddess around 600 CE, and these texts "usually place her on the periphery of Hindu society or on the battlefield." She is often regarded as the Shakti of Shiva, and is closely associated with him in various Puranas. The Kalika Purana depicts her as the "Adi Shakti" (Fundamental Power) and "Para Prakriti" or beyond nature.

 

WORSHIP & MANTRA

Kali could be considered a general concept, like Durga, and is mostly worshiped in the Kali Kula sect of worship. The closest way of direct worship is Maha Kali or Bhadra Kali (Bhadra in Sanskrit means 'gentle'). Kali is worshiped as one of the 10 Mahavidya forms of Adi Parashakti (Goddess Durga) or Bhagavathy according to the region. The mantra for worship is called Devi Argala Stotram.

Sanskrit: सर्वमङ्गलमाङ्गल्ये शिवे सर्वार्थसाधिके । शरण्ये त्र्यम्बके गौरि नारायणि नमोऽस्तु ते ॥

 

ॐ जयंती मंगल काली भद्रकाली कपालिनी । दुर्गा क्षमा शिवा धात्री स्वाहा स्वधा नमोऽस्तु‍ते ॥

(Sarvamaṅgalamāṅgalyē śivē sarvārthasādhikē . śaraṇyē tryambakē gauri nārāyaṇi namō'stu tē.

Oṃ jayantī mangala kālī bhadrakālī kapālinī . durgā kṣamā śivā dhātrī svāhā svadhā namō'stu‍tē.)

 

TANTRA

Goddesses play an important role in the study and practice of Tantra Yoga, and are affirmed to be as central to discerning the nature of reality as are the male deities. Although Parvati is often said to be the recipient and student of Shiva's wisdom in the form of Tantras, it is Kāli who seems to dominate much of the Tantric iconography, texts, and rituals. In many sources Kāli is praised as the highest reality or greatest of all deities. The Nirvana-tantra says the gods Brahma, Vishnu, and Shiva all arise from her like bubbles in the sea, ceaselessly arising and passing away, leaving their original source unchanged. The Niruttara-tantra and the Picchila-tantra declare all of Kāli's mantras to be the greatest and the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra all proclaim Kāli vidyas (manifestations of Mahadevi, or "divinity itself"). They declare her to be an essence of her own form (svarupa) of the Mahadevi.

 

In the Mahanirvana-tantra, Kāli is one of the epithets for the primordial sakti, and in one passage Shiva praises her:

 

At the dissolution of things, it is Kāla [Time] Who will devour all, and by reason of this He is called Mahākāla [an epithet of Lord Shiva], and since Thou devourest Mahākāla Himself, it is Thou who art the Supreme Primordial Kālika. Because Thou devourest Kāla, Thou art Kāli, the original form of all things, and because Thou art the Origin of and devourest all things Thou art called the Adya [the Primordial One]. Re-assuming after Dissolution Thine own form, dark and formless, Thou alone remainest as One ineffable and inconceivable. Though having a form, yet art Thou formless; though Thyself without beginning, multiform by the power of Maya, Thou art the Beginning of all, Creatrix, Protectress, and Destructress that Thou art.

 

The figure of Kāli conveys death, destruction, and the consuming aspects of reality. As such, she is also a "forbidden thing", or even death itself. In the Pancatattva ritual, the sadhaka boldly seeks to confront Kali, and thereby assimilates and transforms her into a vehicle of salvation. This is clear in the work of the Karpuradi-stotra, a short praise of Kāli describing the Pancatattva ritual unto her, performed on cremation grounds. (Samahana-sadhana)

 

He, O Mahākāli who in the cremation-ground, naked, and with dishevelled hair, intently meditates upon Thee and recites Thy mantra, and with each recitation makes offering to Thee of a thousand Akanda flowers with seed, becomes without any effort a Lord of the earth. Oh Kāli, whoever on Tuesday at midnight, having uttered Thy mantra, makes offering even but once with devotion to Thee of a hair of his Shakti [his energy/female companion] in the cremation-ground, becomes a great poet, a Lord of the earth, and ever goes mounted upon an elephant.

 

The Karpuradi-stotra clearly indicates that Kāli is more than a terrible, vicious, slayer of demons who serves Durga or Shiva. Here, she is identified as the supreme mistress of the universe, associated with the five elements. In union with Lord Shiva, she creates and destroys worlds. Her appearance also takes a different turn, befitting her role as ruler of the world and object of meditation. In contrast to her terrible aspects, she takes on hints of a more benign dimension. She is described as young and beautiful, has a gentle smile, and makes gestures with her two right hands to dispel any fear and offer boons. The more positive features exposed offer the distillation of divine wrath into a goddess of salvation, who rids the sadhaka of fear. Here, Kali appears as a symbol of triumph over death.

 

BENGALI TRADITION

Kali is also a central figure in late medieval Bengali devotional literature, with such devotees as Ramprasad Sen (1718–75). With the exception of being associated with Parvati as Shiva's consort, Kāli is rarely pictured in Hindu legends and iconography as a motherly figure until Bengali devotions beginning in the early eighteenth century. Even in Bengāli tradition her appearance and habits change little, if at all.

 

The Tantric approach to Kāli is to display courage by confronting her on cremation grounds in the dead of night, despite her terrible appearance. In contrast, the Bengali devotee appropriates Kāli's teachings adopting the attitude of a child, coming to love her unreservedly. In both cases, the goal of the devotee is to become reconciled with death and to learn acceptance of the way that things are. These themes are well addressed in Rāmprasād's work. Rāmprasād comments in many of his other songs that Kāli is indifferent to his wellbeing, causes him to suffer, brings his worldly desires to nothing and his worldly goods to ruin. He also states that she does not behave like a mother should and that she ignores his pleas:

 

Can mercy be found in the heart of her who was born of the stone? [a reference to Kali as the daughter of Himalaya]

Were she not merciless, would she kick the breast of her lord?

Men call you merciful, but there is no trace of mercy in you, Mother.

You have cut off the heads of the children of others, and these you wear as a garland around your neck.

It matters not how much I call you "Mother, Mother." You hear me, but you will not listen.

 

To be a child of Kāli, Rāmprasād asserts, is to be denied of earthly delights and pleasures. Kāli is said to refrain from giving that which is expected. To the devotee, it is perhaps her very refusal to do so that enables her devotees to reflect on dimensions of themselves and of reality that go beyond the material world.

 

A significant portion of Bengali devotional music features Kāli as its central theme and is known as Shyama Sangeet ("Music of the Night"). Mostly sung by male vocalists, today even women have taken to this form of music. One of the finest singers of Shyāma Sāngeet is Pannalal Bhattacharya.

 

In Bengal, Kāli is venerated in the festival Kali Puja, the new moon day of Ashwin month which coincides with Diwali festival.

 

In a unique form of Kāli worship, Shantipur worships Kāli in the form of a hand painted image of the deity known as Poteshwari (meaning the deity drawn on a piece of cloth).

 

LEGENDS

SLAYER OF RAKTABIJA

In Kāli's most famous legend, Devi Durga (Adi Parashakti) and her assistants, the Matrikas, wound the demon Raktabija, in various ways and with a variety of weapons in an attempt to destroy him. They soon find that they have worsened the situation for with every drop of blood that is dripped from Raktabija he reproduces a clone of himself. The battlefield becomes increasingly filled with his duplicates. Durga, in need of help, summons Kāli to combat the demons. It is said, in some versions, that Goddess Durga actually assumes the form of Goddess Kāli at this time. The Devi Mahatmyam describes:

 

Out of the surface of her (Durga's) forehead, fierce with frown, issued suddenly Kali of terrible countenance, armed with a sword and noose. Bearing the strange khatvanga (skull-topped staff ), decorated with a garland of skulls, clad in a tiger's skin, very appalling owing to her emaciated flesh, with gaping mouth, fearful with her tongue lolling out, having deep reddish eyes, filling the regions of the sky with her roars, falling upon impetuously and slaughtering the great asuras in that army, she devoured those hordes of the foes of the devas.

 

Kali destroys Raktabija by sucking the blood from his body and putting the many Raktabija duplicates in her gaping mouth. Pleased with her victory, Kali then dances on the field of battle, stepping on the corpses of the slain. In the Devi Mahatmya version of this story, Kali is also described as a Matrika and as a Shakti or power of Devi. She is given the epithet Cāṃuṇḍā (Chamunda), i.e. the slayer of the demons Chanda and Munda. Chamunda is very often identified with Kali and is very much like her in appearance and habit.

 

DAKSHINA KALI

In her most famous pose as Daksinakali, popular legends say that Kali, becoming drunk on the blood of her victims on the battlefield, dances with destructive frenzy. She is about to destroy the whole universe when, urged by all the gods, Shiva lies in her way to stop her. In her fury, she fails to see the body of Shiva lying amongst the corpses on the battlefield and steps upon his chest. Realizing Shiva lies beneath her feet, her anger is pacified and she calms her fury. Though not included in any of the puranas, popular legends state that Kali was ashamed at the prospect of keeping her husband beneath her feet and thus stuck her tongue out in shame. The Devi-Bhagavata Purana, which goes into great depths about the goddess Kali, reveals the tongue's actual symbolism.

 

The characteristic icons that depict Kali are the following; unbridled matted hair, open blood shot eyes, open mouth and a drooping tongue; in her hands, she holds a Khadga (bent sword or scimitar) and a human head; she has a girdle of human hands across her waist and an enchanted Shiva lies beneath her feet. Each of these icons represent a deep philosophical epithet. The drooping out-stuck tongue represents her blood-thirst. Lord Shiva beneath her feet represents matter, as Kali is undoubtedly the primeval energy. The depiction of Kali on Shiva shows that without energy, matter lies "dead". This concept has been simplified to a folk-tale depicting a wife placing her foot on her husband and sticking her tongue out in shame. In tantric contexts, the tongue is seen to denote the element (guna) of rajas (energy and action) controlled by sattva.

 

If Kali steps on Shiva with her right foot and holds the sword in her left hand, she is considered to be Dakshina Kali. The Dakshina Kali Temple has important religious associations with the Jagannath Temple and it is believed that Daksinakali is the guardian of the kitchen of the Lord Jagannath Temple. Puranic tradition says that in Puri, Lord Jagannath is regarded as Daksinakalika. Goddess Dakshinakali plays an important role in the 'Niti' of Saptapuri Amavasya.

 

One South Indian tradition tells of a dance contest between Shiva and Kali. After defeating the two demons Sumbha and Nisumbha, Kali takes up residence in the forest of Thiruvalankadu or Thiruvalangadu. She terrorizes the surrounding area with her fierce, disruptive nature. One of Shiva's devotees becomes distracted while performing austerities, and asks Shiva to rid the forest of the destructive goddess. When Shiva arrives, Kali threatens him, claiming the territory as her own. Shiva challenges Kali to a dance contest; both of them dance and Kali matches Shiva in every step that he takes until Shiva takes the "Urdhvatandava" step, by vertically raising his right leg. Kali refuses to perform this step, which would not befit her as a woman, and became pacified.

 

SMASHAN KALI

If the Kali steps out with the left foot and holds the sword in her right hand, she is the terrible form of Mother, the Smashan Kali of the cremation ground. She is worshiped by tantrics, the followers of Tantra, who believe that one's spiritual discipline practiced in a smashan (cremation ground) brings success quickly. Sarda Devi, the consort of Ramakrishna Paramhansa, worshipped Smashan Kali at Dakshineshwar.

 

MATERNAL KALI

Another legend depicts the infant Shiva calming Kali. In this similar story, Kali has defeated her enemies on the battlefield and begun to dance out of control, drunk on the blood of the slain. To calm her down and to protect the stability of the world, Shiva is sent to the battlefield, as an infant, crying aloud. Seeing the child's distress, Kali ceases dancing to care for the helpless infant. She picks him up, kisses his head, and proceeds to breast feed the infant Shiva. This legend is notable because it shows Kali in her benevolent, maternal aspect, with which she is not usually identified.

 

MAHAKALI

Mahakali (Sanskrit: Mahākālī, Devanagari: महाकाली), literally translated as Great Kali, is sometimes considered as a greater form of Kali, identified with the Ultimate reality of Brahman. It can also be used as an honorific of the Goddess Kali, signifying her greatness by the prefix "Mahā-". Mahakali, in Sanskrit, is etymologically the feminized variant of Mahakala or Great Time (which is interpreted also as Death), an epithet of the God Shiva in Hinduism. Mahakali is the presiding Goddess of the first episode of the Devi Mahatmya. Here she is depicted as Devi in her universal form as Shakti. Here Devi serves as the agent who allows the cosmic order to be restored.

 

Kali is depicted in the Mahakali form as having ten heads, ten arms, and ten legs. Each of her ten hands is carrying a various implement which vary in different accounts, but each of these represent the power of one of the Devas or Hindu Gods and are often the identifying weapon or ritual item of a given Deva. The implication is that Mahakali subsumes and is responsible for the powers that these deities possess and this is in line with the interpretation that Mahakali is identical with Brahman. While not displaying ten heads, an "ekamukhi" or one headed image may be displayed with ten arms, signifying the same concept: the powers of the various Gods come only through Her grace.

 

ICONOGRAPHY

Kali is portrayed mostly in two forms: the popular four-armed form and the ten-armed Mahakali form. In both of her forms, she is described as being black in color but is most often depicted as blue in popular Indian art. Her eyes are described as red with intoxication, and in absolute rage, her hair is shown disheveled, small fangs sometimes protrude out of her mouth, and her tongue is lolling. She is often shown naked or just wearing a skirt made of human arms and a garland of human heads. She is also accompanied by serpents and a jackal while standing on a seemingly dead Shiva, usually right foot forward to symbolize the more popular Dakshinamarga or right-handed path, as opposed to the more infamous and transgressive Vamamarga or left-handed path.

 

In the ten-armed form of Mahakali she is depicted as shining like a blue stone. She has ten faces and ten feet and three eyes. She has ornaments decked on all her limbs. There is no association with Shiva.

 

The Kalika Purana describes Kali as possessing a soothing dark complexion, as perfectly beautiful, riding a lion, four-armed, holding a sword and blue lotuses, her hair unrestrained, body firm and youthful.

 

In spite of her seemingly terrible form, Kali Ma is often considered the kindest and most loving of all the Hindu goddesses, as she is regarded by her devotees as the Mother of the whole Universe. And because of her terrible form, she is also often seen as a great protector. When the Bengali saint Ramakrishna once asked a devotee why one would prefer to worship Mother over him, this devotee rhetorically replied, "Maharaj, when they are in trouble your devotees come running to you. But, where do you run when you are in trouble?"

 

According to Ramakrishna, darkness is the Ultimate Mother, or Kali:

 

My Mother is the principle of consciousness. She is Akhanda Satchidananda; indivisible Reality, Awareness, and Bliss. The night sky between the stars is perfectly black. The waters of the ocean depths are the same; The infinite is always mysteriously dark. This inebriating darkness is my beloved Kali.

 

SRI RAMAKRISHNA

This is clear in the works of such contemporary artists as Charles Wish, and Tyeb Mehta, who sometimes take great liberties with the traditional, accepted symbolism, but still demonstrate a true reverence for the Shakta sect.

 

POPULAR FORM

Classic depictions of Kali share several features, as follows:

 

Kali's most common four armed iconographic image shows each hand carrying variously a sword, a trishul (trident), a severed head and a bowl or skull-cup (kapala) catching the blood of the severed head.

 

Two of these hands (usually the left) are holding a sword and a severed head. The Sword signifies Divine Knowledge and the Human Head signifies human Ego which must be slain by Divine Knowledge in order to attain Moksha. The other two hands (usually the right) are in the abhaya (fearlessness) and varada (blessing) mudras, which means her initiated devotees (or anyone worshipping her with a true heart) will be saved as she will guide them here and in the hereafter.

 

She has a garland consisting of human heads, variously enumerated at 108 (an auspicious number in Hinduism and the number of countable beads on a Japa Mala or rosary for repetition of Mantras) or 51, which represents Varnamala or the Garland of letters of the Sanskrit alphabet, Devanagari. Hindus believe Sanskrit is a language of dynamism, and each of these letters represents a form of energy, or a form of Kali. Therefore she is generally seen as the mother of language, and all mantras.

 

She is often depicted naked which symbolizes her being beyond the covering of Maya since she is pure (nirguna) being-consciousness-bliss and far above prakriti. She is shown as very dark as she is brahman in its supreme unmanifest state. She has no permanent qualities - she will continue to exist even when the universe ends. It is therefore believed that the concepts of color, light, good, bad do not apply to her - she is the pure, un-manifested energy, the Adi-shakti.

 

SHIVA IN KALI ICONOGRAPHY

In both these images she is shown standing on the prone, inert or dead body of Shiva. There is a legend for the reason behind her standing on what appears to be Shiva's corpse, which translates as follows:

 

Once Kali had destroyed all the demons in battle, she began a terrific dance out of the sheer joy of victory. All the worlds or lokas began to tremble and sway under the impact of her dance. So, at the request of all the Gods, Shiva himself asked her to desist from this behavior. However, she was too intoxicated to listen. Hence, Shiva lay like a corpse among the slain demons in order to absorb the shock of the dance into himself. When Kali eventually stepped upon Shiva, she realized she was trampling and hurting her husband and bit her tongue in shame.

 

The story described here is a popular folk tale and not described or hinted in any of the puranas. The puranic interpretation is as follows:

 

Once, Parvati asks Shiva to chose the one form among her 10 forms which he likes most. To her surprise, Shiva reveals that he is most comfortable with her Kali form, in which she is bereft of her jewellery, her human-form, her clothes, her emotions and where she is only raw, chaotic energy, where she is as terrible as time itself and even greater than time. As Parvati takes the form of Kali, Shiva lies at her feet and requests her to place her foot on his chest, upon his heart. Once in this form, Shiva requests her to have this place, below her feet in her iconic image which would be worshiped throughout.

 

This idea has been explored in the Devi-Bhagavata Purana and is most popular in the Shyama Sangeet, devotional songs to Kali from the 12th to 15th centuries.

 

The Tantric interpretation of Kali standing on top of her husband is as follows:

 

The Shiv tattava (Divine Consciousness as Shiva) is inactive, while the Shakti tattava (Divine Energy as Kali) is active. Shiva and Kali represent Brahman, the Absolute pure consciousness which is beyond all names, forms and activities. Kali, on the other hand, represents the potential (and manifested) energy responsible for all names, forms and activities. She is his Shakti, or creative power, and is seen as the substance behind the entire content of all consciousness. She can never exist apart from Shiva or act independently of him, just as Shiva remains a mere corpse without Kali i.e., Shakti, all the matter/energy of the universe, is not distinct from Shiva, or Brahman, but is rather the dynamic power of Brahman. Hence, Kali is Para Brahman in the feminine and dynamic aspect while Shiva is the male aspect and static. She stands as the absolute basis for all life, energy and beneath her feet lies, Shiva, a metaphor for mass, which cannot retain its form without energy.

 

While this is an advanced concept in monistic Shaktism, it also agrees with the Nondual Trika philosophy of Kashmir, popularly known as Kashmir Shaivism and associated most famously with Abhinavagupta. There is a colloquial saying that "Shiva without Shakti is Shava" which means that without the power of action (Shakti) that is Mahakali (represented as the short "i" in Devanagari) Shiva (or consciousness itself) is inactive; Shava means corpse in Sanskrit and the play on words is that all Sanskrit consonants are assumed to be followed by a short letter "a" unless otherwise noted. The short letter "i" represents the female power or Shakti that activates Creation. This is often the explanation for why She is standing on Shiva, who is either Her husband and complement in Shaktism or the Supreme Godhead in Shaivism.

 

To properly understand this complex Tantric symbolism it is important to remember that the meaning behind Shiva and Kali does not stray from the non-dualistic parlance of Shankara or the Upanisads. According to both the Mahanirvana and Kularnava Tantras, there are two distinct ways of perceiving the same absolute reality. The first is a transcendental plane which is often described as static, yet infinite. It is here that there is no matter, there is no universe and only consciousness exists. This form of reality is known as Shiva, the absolute Sat-Chit-Ananda - existence, knowledge and bliss. The second is an active plane, an immanent plane, the plane of matter, of Maya, i.e., where the illusion of space-time and the appearance of an actual universe does exist. This form of reality is known as Kali or Shakti, and (in its entirety) is still specified as the same Absolute Sat-Chit-Ananda. It is here in this second plane that the universe (as we commonly know it) is experienced and is described by the Tantric seer as the play of Shakti, or God as Mother Kali.

 

From a Tantric perspective, when one meditates on reality at rest, as absolute pure consciousness (without the activities of creation, preservation or dissolution) one refers to this as Shiva or Brahman. When one meditates on reality as dynamic and creative, as the Absolute content of pure consciousness (with all the activities of creation, preservation or dissolution) one refers to it as Kali or Shakti. However, in either case the yogini or yogi is interested in one and the same reality - the only difference being in name and fluctuating aspects of appearance. It is this which is generally accepted as the meaning of Kali standing on the chest of Shiva.

 

Although there is often controversy surrounding the images of divine copulation, the general consensus is benign and free from any carnal impurities in its substance. In Tantra the human body is a symbol for the microcosm of the universe; therefore sexual process is responsible for the creation of the world. Although theoretically Shiva and Kali (or Shakti) are inseparable, like fire and its power to burn, in the case of creation they are often seen as having separate roles. With Shiva as male and Kali as female it is only by their union that creation may transpire. This reminds us of the prakrti and purusa doctrine of Samkhya wherein prakāśa- vimarśa has no practical value, just as without prakrti, purusa is quite inactive. This (once again) stresses the interdependencies of Shiva and Shakti and the vitality of their union.

 

Gopi Krishna proposed that Kali standing on the dead Shiva or Shava (Sanskrit for dead body) symbolised the helplessness of a person undergoing the changing process (psychologically and physiologically) in the body conducted by the Kundalini Shakti.

 

DEVELOPMENT

In the later traditions, Kali has become inextricably linked with Shiva. The unleashed form of Kali often becomes wild and uncontrollable, and only Shiva is able to tame her just as only Kali can tame Shiva. This is both because she is often a transformed version of one of his consorts and because he is able to match her wildness.

 

The ancient text of Kali Kautuvam describes her competition with Shiva in dance, from which the sacred 108 Karanas appeared. Shiva won the competition by acting the urdva tandava, one of the Karanas, by raising his feet to his head. Other texts describe Shiva appearing as a crying infant and appealing to her maternal instincts. While Shiva is said to be able to tame her, the iconography often presents her dancing on his fallen body, and there are accounts of the two of them dancing together, and driving each other to such wildness that the world comes close to unravelling.

 

Shiva's involvement with Tantra and Kali's dark nature have led to her becoming an important Tantric figure. To the Tantric worshippers, it was essential to face her Curse, the terror of death, as willingly as they accepted Blessings from her beautiful, nurturing, maternal aspect. For them, wisdom meant learning that no coin has only one side: as death cannot exist without life, so life cannot exist without death. Kali's role sometimes grew beyond that of a chaos - which could be confronted - to that of one who could bring wisdom, and she is given great metaphysical significance by some Tantric texts. The Nirvāna-tantra clearly presents her uncontrolled nature as the Ultimate Reality, claiming that the trimurti of Brahma, Vishnu and Rudra arise and disappear from her like bubbles from the sea. Although this is an extreme case, the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra declare her the svarupa (own-being) of the Mahadevi (the great Goddess, who is in this case seen as the combination of all devis).

 

The final stage of development is the worshipping of Kali as the Great Mother, devoid of her usual violence. This practice is a break from the more traditional depictions. The pioneers of this tradition are the 18th century Shakta poets such as Ramprasad Sen, who show an awareness of Kali's ambivalent nature. Ramakrishna, the 19th century Bengali saint, was also a great devotee of Kali; the western popularity of whom may have contributed to the more modern, equivocal interpretations of this Goddess. Rachel McDermott's work, however, suggests that for the common, modern worshipper, Kali is not seen as fearful, and only those educated in old traditions see her as having a wrathful component. Some credit to the development of Devi must also be given to Samkhya. Commonly referred to as the Devi of delusion, Mahamaya or Durga, acting in the confines of (but not being bound by) the nature of the three gunas, takes three forms: Maha-Kali, Maha-Lakshmi and Maha-Saraswati, being her tamas-ika, rajas-ika and sattva-ika forms. In this sense, Kali is simply part of a larger whole.

 

Like Sir John Woodroffe and Georg Feuerstein, many Tantric scholars (as well as sincere practitioners) agree that, no matter how propitious or appalling you describe them, Shiva and Devi are simply recognizable symbols for everyday, abstract (yet tangible) concepts such as perception, knowledge, space-time, causation and the process of liberating oneself from the confines of such things. Shiva, symbolizing pure, absolute consciousness, and Devi, symbolizing the entire content of that consciousness, are ultimately one and the same - totality incarnate, a micro-macro-cosmic amalgamation of all subjects, all objects and all phenomenal relations between the "two." Like man and woman who both share many common, human traits yet at the same time they are still different and, therefore, may also be seen as complementary.

 

Worshippers prescribe various benign and horrific qualities to Devi simply out of practicality. They do this so they may have a variety of symbols to choose from, symbols which they can identify and relate with from the perspective of their own, ever-changing time, place and personal level of unfolding. Just like modern chemists or physicists use a variety of molecular and atomic models to describe what is unperceivable through rudimentary, sensory input, the scientists of ontology and epistemology must do the same. One of the underlying distinctions of Tantra, in comparison to other religions, is that it allows the devotee the liberty to choose from a vast array of complementary symbols and rhetoric which suit one's evolving needs and tastes. From an aesthetic standpoint, nothing is interdict and nothing is orthodox. In this sense, the projection of some of Devi's more gentle qualities onto Kali is not sacrilege and the development of Kali really lies in the practitioner, not the murthi.

 

A TIME magazine article of October 27, 1947, used Kali as a symbol and metaphor for the human suffering in British India during its partition that year.

 

Swami Vivekananda wrote his favorite poem Kali the Mother in 1898.

 

IN NEW AGE & NEOPAGANISM

An academic study of Western Kali enthusiasts noted that, "as shown in the histories of all cross-cultural religious transplants, Kali devotionalism in the West must take on its own indigenous forms if it is to adapt to its new environment." The adoption of Kali by the West has raised accusations of cultural appropriation:

 

A variety of writers and thinkers have found Kali an exciting figure for reflection and exploration, notably feminists and participants in New Age spirituality who are attracted to goddess worship. Kali is a symbol of wholeness and healing, associated especially with repressed female power and sexuality. [However, such interpretations often exhibit] confusion and misrepresentation, stemming from a lack of knowledge of Hindu history among these authors, draw upon materials written by scholars of the Hindu religious tradition. The majority instead rely chiefly on other popular feminist sources, almost none of which base their interpretations on a close reading of Kali's Indian background. The most important issue arising from this discussion - even more important than the question of 'correct' interpretation - concerns the adoption of other people's religious symbols. It is hard to import the worship of a goddess from another culture: religious associations and connotations have to be learned, imagined or intuited when the deep symbolic meanings embedded in the native culture are not available.

 

WIKIPEDIA

www.charliecolmer.com

Mary Hartigan's Restaurant located at 910 Washington St. in Dedham,MA. The restaurant was demolished in 2011 and by 2014 a medical office had been constructed on the site. The restaurant was constructed as a Dutchland Farms Restaurant in the early 1930s although by the mid 1940s was renamed after its proprietor Mary Hartignan. The Hartigan family operated the restaurant until the early 1990s. Once the Hartigan's sold the property the building was converted into an interdependently owned restaurant which operated under the name Piccadilly Pub. The Piccadilly Pub restaurant operated for a few years before being converted into a seafood restaurant which closed in 2010. Note the building had a very similar design to the highly successful (at the time) Howard Johnson's Restaurants complete with tiled roof and cupola.

Elephants may be famed for their wrinkly exterior, but when it comes to Gillian Higgins' sculpture, it really is what’s on the inside that counts. This design, created by anatomist Gillian, depicts the intricate, interdependent and wonder of the internal systems on which life depends. Gillian believes that having a deeper understanding of our inner workings is essential in helping us to make wise decisions about our health and improve personal well being.

 

Designed by: Gillian Higgins

Gillian Higgins is the director of Horses Inside Out, an organisation which shows how understanding anatomy, physiology and bio mechanics can improve well being, performance and reduce risk of injury. Gillian’s unique trademark is painting the anatomical systems on the sides of live horses which she then uses as live canvases. Initially trained as a remedial sports and massage therapist, Gillian is also a McTimoney animal practitioner, author, British Horse Society Senior Coach and a competitive rider. She is regularly invited to give talks and demonstrations all over the world and has visited Australia, South Africa, North and South America as well as Europe and the British Isles.

 

Sponsored by: B Braun

Auction Price: £6000

 

Summer 2016, a herd of elephant sculptures descended on Sheffield for the biggest public art event the city has ever seen!

58 elephant sculptures, each uniquely decorated by artists, descended on Sheffield’s parks and open spaces, creating one of the biggest mass participation arts events the city has ever seen. Did you find them all?

The trail of elephants celebrates Sheffield’s creativity with over 75% of artists from the city. Some well-known names include Pete McKee, James Green, Jonathan Wilkinson and Lydia Monks – each of which has put their own creative mark on a 1.6m tall fibreglass elephant sculpture. They are all very difference, take a selfie with your favourite as they will be on display until the end of September.

International artist Mark Alexander, who is currently working with Rembrandt for an exhibition in Berlin, flew to Sheffield especially to paint his elephant and international players from the World Snooker Championship signed SnookHerd, an elephant celebrating the heritage of snooker in Sheffield.

The Arctic Monkeys, famous for their love of their home city, added their signatures to their own personalised sculpture which pays homage to the striking sound wave cover of the band’s 2013 album “AM”.

By supporting the Herd of Sheffield you are investing in the future of Sheffield Children’s Hospital. Every penny raised will go towards our Artfelt programme, which transforms the hospital’s walls and spaces with bright art, helping children recover in an environment tailored to them. The programme also puts on workshops for youngsters to provide distraction during anxious moments – such as before an operation, and to breakup long stays on the wards.

This exciting Wild in Art event brought to you by The Children’s Hospital Charity will:

Unite our city – bringing businesses, communities, artists, individuals and schools together to create a FREE sculpture trail which is accessible to all.

Attract more visitors – both nationally and regionally as well as encouraging thousands of people to become a tourist in their own city.

Invest in the future – with a city wide education programme that can be used for years to come and by funding a life-saving piece of medical equipment at Sheffield Children’s Hospital from the Herd auction at the end of the trail.

Showcase our city – celebrating Sheffield’s heritage and cementing our status as a vibrant and culturally exciting city through this world-class initiative.

 

The Herd of Sheffield Farewell Weekend was held on 14-16 October and was your chance to say a last goodbye to all 58 large elephant sculptures as they gather in one place for a final send-off at Meadowhall.

This special event gave visitors a chance to see the entire herd in all its glory – from the signed Arctic Monkeys’ ‘AM’ elephant, right through to ‘SnookHerd’, autographed by a host of international snooker players including current world champion Mark Selby.

Please note that the Little Herd elephants will not be on display as they will be returned to their school for pupils to enjoy.

Meadowhall, along with its joint owners, British Land are very proud to be supporting The Children’s Hospital Charity as host sponsors for the Herd of Sheffield Farewell Weekend.

 

Auction: Hundreds of elephant enthusiasts gathered at the Crucible on 20 October for the Herd of Sheffield Auction, which raised a total of £410,600 for The Children’s Hospital Charity.

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Great Britain, London, Trafalgar Square Fourth Plinth, the Programme is the most spoken about contemporary art prize in the UK. Funded by the Mayor of London & supported by Arts Council England, the programme invites world-class artists to make astonishing works for the centre of the city.

The new commission Gift Horse by artist Hans Haacke, was unveiled on Thursday 5 March 2015 in London’s Trafalgar Square.

The Gift Horse is a bronze skeletal, riderless horse, which has a ribbon wrapped around one leg, displaying the electroluminescent film ticker from the London Stock Exchange, completing the link between power, money & history represented on the fourth plinth

 

Hans Haacke German-American, a conceptual artist, born 1936 in Cologne & studied art in Germany & Philadelphia USA, who lives & works in New York. Haacke’s early work employed physical & organic processes, such as condensation, in what he called ‘systems’, until his focus shifted to the socio-political field of equally interdependent dynamics. For the last four decades, Haacke has been examining relationships between art, power & money; he addressed issues of free expression & public responsibilities in democratic societies.

However the director of the Tate gallery, Nicholas Serota had a very different interpretation, he thinks it's clearly about the fragility of power systems, the state & the financial systems!!

 

...Danke, Xièxie 谢谢, Thanks, Gracias, Merci, Grazie, Obrigado, Arigatô, Dhanyavad, Chokrane to you & over

3,5 million visits in my photostream with countless motivating comments

  

The Global Centre for Plurialism in Lower Town, Ottawa, Ontario, Canada.

 

The Global Centre for Pluralism is an independent, not-for-profit international research and education centre located in Ottawa, Canada. Inspired by the example of Canada’s inclusive approach to citizenship, the Centre works to advance respect for diversity worldwide, believing that openness and understanding toward the cultures, social structures, values and faiths of other peoples are essential to the survival of an interdependent world.

 

For folks familiar with Ottawa, this building previously served as the home of the Public Archives of Canada from 1906 to 1967 and the Canadian War Museum from 1967 to 2005. The building was built from 1904 to 1906 and is designated a National Historic Site of Canada and a Classified Federal Heritage Building.

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

The Dakshinkali Temple is located 22 kilometers from Kathmandu next to the village of Pharping. It's one of the main temples in Nepal. Twice every week thousands of people come here to worship the goddess Kali by sacrificing life animals, particularly cockerels and uncastrated male goats.

 

GODDESS KALI

Kālī (/ˈkɑːli/; Sanskrit: काली & Bengali: কালী; IPA: [kɑːliː]), also known as Kālikā (Sanskrit: कालिका), is the Hindu goddess associated with empowerment, or shakti. She is the fierce aspect of the goddess Durga. The name of Kali means black one and force of time; she is therefore called the Goddess of Time, Change, Power, Creation, Preservation, and Destruction. Her earliest appearance is that of a destroyer principally of evil forces. Various Shakta Hindu cosmologies, as well as Shākta Tantric beliefs, worship her as the ultimate reality or Brahman; and recent devotional movements re-imagine Kāli as a benevolent mother goddess. She is often portrayed standing or dancing on her husband, the god Shiva, who lies calm and prostrate beneath her. Worshipped throughout India but particularly South India, Bengal, and Assam, Kali is both geographically and culturally marginal.

 

ETYMOLOGY

Kālī is the feminine form of kālam ("black, dark coloured"). Kāla primarily means "time", but also means "black"; hence, Kālī means "the black one" or "beyond time". Kāli is strongly associated with Shiva, and Shaivas derive the masculine Kāla (an epithet of Shiva) from her feminine name. A nineteenth-century Sanskrit dictionary, the Shabdakalpadrum, states: कालः शिवः। तस्य पत्नीति - काली। kālaḥ śivaḥ। tasya patnīti kālī - "Shiva is Kāla, thus, his consort is Kāli".

 

Other names include Kālarātri ("black night"), as described above, and Kālikā ("relating to time"), and Kallie ("black alchemist"). Coburn notes that the name Kālī can be used as a proper name, or as a description of color.

 

Kāli's association with darkness stands in contrast to her consort, Shiva, whose body is covered by the white ashes of the cremation ground (Sanskrit: śmaśāna) where he meditates, and with which Kāli is also associated, as śmaśāna-kālī.

 

ORIGINS

Hugh Urban notes that although the word Kālī appears as early as the Atharva Veda, the first use of it as a proper name is in the Kathaka Grhya Sutra (19.7). Kali is the name of one of the seven tongues of Agni, the [Rigvedic] God of Fire, in the Mundaka Upanishad (2:4), but it is unlikely that this refers to the goddess. The first appearance of Kāli in her present form is in the Sauptika Parvan of the Mahabharata (10.8.64). She is called Kālarātri (literally, "black night") and appears to the Pandava soldiers in dreams, until finally she appears amidst the fighting during an attack by Drona's son Ashwatthama. She most famously appears in the sixth century Devi Mahatmyam as one of the shaktis of Mahadevi, and defeats the demon Raktabija ("Bloodseed"). The tenth-century Kalika Purana venerates Kāli as the ultimate reality.

 

According to David Kinsley, Kāli is first mentioned in Hinduism as a distinct goddess around 600 CE, and these texts "usually place her on the periphery of Hindu society or on the battlefield." She is often regarded as the Shakti of Shiva, and is closely associated with him in various Puranas. The Kalika Purana depicts her as the "Adi Shakti" (Fundamental Power) and "Para Prakriti" or beyond nature.

 

WORSHIP AND MANTRA

Kali could be considered a general concept, like Durga, and is mostly worshiped in the Kali Kula sect of worship. The closest way of direct worship is Maha Kali or Bhadra Kali (Bhadra in Sanskrit means 'gentle'). Kali is worshiped as one of the 10 Mahavidya forms of Adi Parashakti (Goddess Durga) or Bhagavathy according to the region. The mantra for worship is

 

Sanskrit: सर्वमङ्गलमाङ्गल्ये शिवे सर्वार्थसाधिके । शरण्ये त्र्यम्बके गौरि नारायणि नमोऽस्तु ते ॥

 

ॐ जयंती मंगल काली भद्रकाली कपालिनी । दुर्गा शिवा क्षमा धात्री स्वाहा स्वधा नमोऽस्तु‍ते ॥

 

(Sarvamaṅgalamāṅgalyē śivē sarvārthasādhikē . śaraṇyē tryambakē gauri nārāyaṇi namō'stu tē.

 

Oṃ jayantī mangala kālī bhadrakālī kapālinī . durgā śivā ksamā dhātrī svāhā svadhā namō'stu‍tē.)

 

YANTRA

Goddesses play an important role in the study and practice of Tantra Yoga, and are affirmed to be as central to discerning the nature of reality as are the male deities. Although Parvati is often said to be the recipient and student of Shiva's wisdom in the form of Tantras, it is Kali who seems to dominate much of the Tantric iconography, texts, and rituals. In many sources Kāli is praised as the highest reality or greatest of all deities. The Nirvana-tantra says the gods Brahma, Vishnu, and Shiva all arise from her like bubbles in the sea, ceaselessly arising and passing away, leaving their original source unchanged. The Niruttara-tantra and the Picchila-tantra declare all of Kāli's mantras to be the greatest and the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra all proclaim Kāli vidyas (manifestations of Mahadevi, or "divinity itself"). They declare her to be an essence of her own form (svarupa) of the Mahadevi.In the Mahanirvana-tantra, Kāli is one of the epithets for the primordial sakti, and in one passage Shiva praises her:At the dissolution of things, it is Kāla [Time]. Who will devour all, and by reason of this He is called Mahākāla [an epithet of Lord Shiva], and since Thou devourest Mahākāla Himself, it is Thou who art the Supreme Primordial Kālika. Because Thou devourest Kāla, Thou art Kāli, the original form of all things, and because Thou art the Origin of and devourest all things Thou art called the Adya [the Primordial One]. Re-assuming after Dissolution Thine own form, dark and formless, Thou alone remainest as One ineffable and inconceivable. Though having a form, yet art Thou formless; though Thyself without beginning, multiform by the power of Maya, Thou art the Beginning of all, Creatrix, Protectress, and Destructress that Thou art. The figure of Kāli conveys death, destruction, and the consuming aspects of reality. As such, she is also a "forbidden thing", or even death itself. In the Pancatattva ritual, the sadhaka boldly seeks to confront Kali, and thereby assimilates and transforms her into a vehicle of salvation. This is clear in the work of the Karpuradi-stotra, a short praise of Kāli describing the Pancatattva ritual unto her, performed on cremation grounds. (Samahana-sadhana)He, O Mahākāli who in the cremation-ground, naked, and with dishevelled hair, intently meditates upon Thee and recites Thy mantra, and with each recitation makes offering to Thee of a thousand Akanda flowers with seed, becomes without any effort a Lord of the earth. Oh Kāli, whoever on Tuesday at midnight, having uttered Thy mantra, makes offering even but once with devotion to Thee of a hair of his Shakti [his energy/female companion] in the cremation-ground, becomes a great poet, a Lord of the earth, and ever goes mounted upon an elephant.The Karpuradi-stotra clearly indicates that Kāli is more than a terrible, vicious, slayer of demons who serves Durga or Shiva. Here, she is identified as the supreme mistress of the universe, associated with the five elements. In union with Lord Shiva, she creates and destroys worlds. Her appearance also takes a different turn, befitting her role as ruler of the world and object of meditation. In contrast to her terrible aspects, she takes on hints of a more benign dimension. She is described as young and beautiful, has a gentle smile, and makes gestures with her two right hands to dispel any fear and offer boons. The more positive features exposed offer the distillation of divine wrath into a goddess of salvation, who rids the sadhaka of fear. Here, Kali appears as a symbol of triumph over death.

 

BENGALI TRADITION

Kali is also a central figure in late medieval Bengali devotional literature, with such devotees as Ramprasad Sen (1718–75). With the exception of being associated with Parvati as Shiva's consort, Kāli is rarely pictured in Hindu legends and iconography as a motherly figure until Bengali devotions beginning in the early eighteenth century. Even in Bengāli tradition her appearance and habits change little, if at all.

 

The Tantric approach to Kāli is to display courage by confronting her on cremation grounds in the dead of night, despite her terrible appearance. In contrast, the Bengali devotee appropriates Kāli's teachings adopting the attitude of a child, coming to love her unreservedly. In both cases, the goal of the devotee is to become reconciled with death and to learn acceptance of the way that things are. These themes are well addressed in Rāmprasād's work. Rāmprasād comments in many of his other songs that Kāli is indifferent to his wellbeing, causes him to suffer, brings his worldly desires to nothing and his worldly goods to ruin. He also states that she does not behave like a mother should and that she ignores his pleas:

 

Can mercy be found in the heart of her who was born of the stone? [a reference to Kali as the daughter of Himalaya]

Were she not merciless, would she kick the breast of her lord?

Men call you merciful, but there is no trace of mercy in you, Mother.

You have cut off the heads of the children of others, and these you wear as a garland around your neck.

It matters not how much I call you "Mother, Mother." You hear me, but you will not listen.

 

To be a child of Kāli, Rāmprasād asserts, is to be denied of earthly delights and pleasures. Kāli is said to refrain from giving that which is expected. To the devotee, it is perhaps her very refusal to do so that enables her devotees to reflect on dimensions of themselves and of reality that go beyond the material world.

 

A significant portion of Bengali devotional music features Kāli as its central theme and is known as Shyama Sangeet ("Music of the Night"). Mostly sung by male vocalists, today even women have taken to this form of music. One of the finest singers of Shyāma Sāngeet is Pannalal Bhattacharya.

 

In Bengal, Kāli is venerated in the festival Kali Puja, the new moon day of Ashwin month which coincides with Diwali festival.

 

In a unique form of Kāli worship, Shantipur worships Kāli in the form of a hand painted image of the deity known as Poteshwari (meaning the deity drawn on a piece of cloth).

 

LEGENDS

SLAYER AND RAKTABIJA

In Kāli's most famous legend, Devi Durga (Adi Parashakti) and her assistants, the Matrikas, wound the demon Raktabija, in various ways and with a variety of weapons in an attempt to destroy him. They soon find that they have worsened the situation for with every drop of blood that is dripped from Raktabija he reproduces a clone of himself. The battlefield becomes increasingly filled with his duplicates. Durga, in need of help, summons Kāli to combat the demons. It is said, in some versions, that Goddess Durga actually assumes the form of Goddess Kāli at this time. The Devi Mahatmyam describes:

 

Out of the surface of her (Durga's) forehead, fierce with frown, issued suddenly Kali of terrible countenance, armed with a sword and noose. Bearing the strange khatvanga (skull-topped staff ), decorated with a garland of skulls, clad in a tiger's skin, very appalling owing to her emaciated flesh, with gaping mouth, fearful with her tongue lolling out, having deep reddish eyes, filling the regions of the sky with her roars, falling upon impetuously and slaughtering the great asuras in that army, she devoured those hordes of the foes of the devas.

 

Kali consumes Raktabija and his duplicates, and dances on the corpses of the slain. In the Devi Mahatmya version of this story, Kali is also described as a Matrika and as a Shakti or power of Devi. She is given the epithet Cāṃuṇḍā (Chamunda), i.e. the slayer of the demons Chanda and Munda. Chamunda is very often identified with Kali and is very much like her in appearance and habit.

  

DAKSHINA KALI

In her most famous pose as Daksinakali, popular legends say that Kali, drunk on the blood of her victims, is about to destroy the whole universe when, urged by all the gods, Shiva lies in her way to stop her, and she steps upon his chest. Recognizing Shiva beneath her feet, she calms herself. Though not included in any of the puranas, popular legends state that Kali was ashamed at the prospect of keeping her husband beneath her feet and thus stuck her tongue out in shame. The Devi-Bhagavata Purana, which goes into great depths about the goddess Kali, reveals the tongue's actual symbolism.

 

The characteristic icons that depict Kali are the following; unbridled matted hair, open blood shot eyes, open mouth and a drooping tongue; in her hands, she holds a Khadga (bent sword or scimitar) and a human head; she has a girdle of human hands across her waist, and Shiva lies beneath her feet. The drooping out-stuck tongue represents her blood-thirst. Lord Shiva beneath her feet represents matter, as Kali energy. The depiction of Kali on Shiva shows that without energy, matter lies "dead". This concept has been simplified to a folk-tale depicting a wife placing her foot

 

on her husband and sticking her tongue out in shame. In tantric contexts, the tongue is seen to denote the element (guna) of rajas (energy and action) controlled by sattva.

 

If Kali steps on Shiva with her right foot and holds the sword in her left hand, she is considered to be Dakshina Kali. The Dakshina Kali Temple has important religious associations with the Jagannath Temple and it is believed that Daksinakali is the guardian of the kitchen of the Lord Jagannath Temple. Puranic tradition says that in Puri, Lord Jagannath is regarded as Daksinakalika. Goddess Dakshinakali plays an important role in the 'Niti' of Saptapuri Amavasya.

 

One South Indian tradition tells of a dance contest between Shiva and Kali. After defeating the two demons Sumbha and Nisumbha, Kali takes up residence in the forest of Thiruvalankadu or Thiruvalangadu. She terrorizes the surrounding area with her fierce, disruptive nature. One of Shiva's devotees becomes distracted while performing austerities, and asks Shiva to rid the forest of the destructive goddess. When Shiva arrives, Kali threatens him, and Shiva challenges Kali to a dance contest, wherein Kali matches Shiva until Shiva takes the "Urdhvatandava" step, vertically raising his right leg. Kali refuses to perform this step, which would not befit her as a woman, and becomes pacified.

 

SMASHAN KALI

If the Kali steps out with the left foot and holds the sword in her right hand, she is the terrible form of Mother, the Smashan Kali of the cremation ground. She is worshiped by tantrics, the followers of Tantra, who believe that one's spiritual discipline practiced in a smashan (cremation ground) brings success quickly. Sarda Devi, the consort of Ramakrishna Paramhansa, worshipped Smashan Kali at Dakshineshwar.

 

MATERNAL KALI

At the time of samundra manthan when amrit came out, along with that came out poison which was going to destroy the world hence on the request of all the gods, Lord Shiva drank it to save the world but as he is beyond death he didn't die but was very much in pain due to the poison effect hence he became a child so that Kali can feed him with her milk which will sooth out the poison effect.

 

MAHAKALI

Mahakali (Sanskrit: Mahākālī, Devanagari: महाकाली), literally translated as Great Kali, is sometimes considered as a greater form of Kali, identified with the Ultimate reality of Brahman. It can also be used as an honorific of the Goddess Kali, signifying her greatness by the prefix "Mahā-". Mahakali, in Sanskrit, is etymologically the feminized variant of Mahakala or Great Time (which is interpreted also as Death), an epithet of the God Shiva in Hinduism. Mahakali is the presiding Goddess of the first episode of the Devi Mahatmya. Here she is depicted as Devi in her universal form as Shakti. Here Devi serves as the agent who allows the cosmic order to be restored.

 

Kali is depicted in the Mahakali form as having ten heads, ten arms, and ten legs. Each of her ten hands is carrying a various implement which vary in different accounts, but each of these represent the power of one of the Devas or Hindu Gods and are often the identifying weapon or ritual item of a given Deva. The implication is that Mahakali subsumes and is responsible for the powers that these deities possess and this is in line with the interpretation that Mahakali is identical with Brahman. While not displaying ten heads, an "ekamukhi" or one headed image may be displayed with ten arms, signifying the same concept: the powers of the various Gods come only through Her grace.

 

ICONOGRAPHY

Kali is portrayed mostly in two forms: the popular four-armed form and the ten-armed Mahakali form. In both of her forms, she is described as being black in color but is most often depicted as blue in popular Indian art. Her eyes are described as red with intoxication, and in absolute rage, her hair is shown disheveled, small fangs sometimes protrude out of her mouth, and her tongue is lolling. She is often shown naked or just wearing a skirt made of human arms and a garland of human heads. She is also accompanied by serpents and a jackal while standing on a seemingly dead Shiva, usually right foot forward to symbolize the more popular Dakshinamarga or right-handed path, as opposed to the more infamous and transgressive Vamamarga or left-handed path.

 

In the ten-armed form of Mahakali she is depicted as shining like a blue stone. She has ten faces, ten feet, and three eyes for each head. She has ornaments decked on all her limbs. There is no association with Shiva.

 

The Kalika Purana describes Kali as possessing a soothing dark complexion, as perfectly beautiful, riding a lion, four-armed, holding a sword and blue lotuses, her hair unrestrained, body firm and youthful.

 

In spite of her seemingly terrible form, Kali Ma is often considered the kindest and most loving of all the Hindu goddesses, as she is regarded by her devotees as the Mother of the whole Universe. And because of her terrible form, she is also often seen as a great protector. When the Bengali saint Ramakrishna once asked a devotee why one would prefer to worship Mother over him, this devotee rhetorically replied, "Maharaj", when they are in trouble your devotees come running to you. But, where do you run when you are in trouble?"

 

According to Ramakrishna, darkness is the Ultimate Mother, or Kali:

 

My Mother is the principle of consciousness. She is Akhanda Satchidananda;

indivisible Reality, Awareness, and Bliss. The night sky between the stars is perfectly black.

The waters of the ocean depths are the same; The infinite is always mysteriously dark.

This inebriating darkness is my beloved Kali.

—Sri Ramakrishna

This is clear in the works of such contemporary artists as Charles Wish, and Tyeb Mehta, who sometimes take great liberties with the traditional, accepted symbolism, but still demonstrate a true reverence for the Shakta sect.

 

POPULAR FORM

Classic depictions of Kali share several features, as follows:

 

Kali's most common four armed iconographic image shows each hand carrying variously a sword, a trishul (trident), a severed head, and a bowl or skull-cup (kapala) catching the blood of the severed head.

 

Two of these hands (usually the left) are holding a sword and a severed head. The Sword signifies Divine Knowledge and the Human Head signifies human Ego which must be slain by Divine Knowledge in order to attain Moksha. The other two hands (usually the right) are in the abhaya (fearlessness) and varada (blessing) mudras, which means her initiated devotees (or anyone worshipping her with a true heart) will be saved as she will guide them here and in the hereafter.

 

She has a garland consisting of human heads, variously enumerated at 108 (an auspicious number in Hinduism and the number of countable beads on a Japa Mala or rosary for repetition of Mantras) or 51, which represents Varnamala or the Garland of letters of the Sanskrit alphabet, Devanagari. Hindus believe Sanskrit is a language of dynamism, and each of these letters represents a form of energy, or a form of Kali. Therefore, she is generally seen as the mother of language, and all mantras.

 

She is often depicted naked which symbolizes her being beyond the covering of Maya since she is pure (nirguna) being-consciousness-bliss and far above prakriti. She is shown as very dark as she is brahman in its supreme unmanifest state. She has no permanent qualities - she will continue to exist even when the universe ends. It is therefore believed that the concepts of color, light, good, bad do not apply to her - she is the pure, un-manifested energy, the Adi-shakti.

 

Kali as the Symbol of Creation , Freedom , Preservation and Destruction

 

The head that hangs in Kali's hand is a symbol of Ego and the scimitar which she is holding represents power and energy.It is believed that Kali is protecting the human race by that scimitar and also destroying the negativity and ego within human being. The body lying under Kali symbolizes ruination, is actually a form of Shiva. Kali steps her leg on the chest of the body and suppress ruination . Since she is standing on the pure white chest of Lord Shiva who, as pure primal awareness, lays in a passive reclining position, peacefully lies with his eyes half open in a state of bliss. Her hair is long, black and flowing freely depicting Her freedom from convention and the confines of conceptualization. The white teeth which Kali has stands for conscience and her red tongue represents greed. By pressing her white teeth on her tongue Kali refers to control greed.The goddess may appear terrible from outside but every symbol in Kali signifies truth of life. Since the earth was created out of darkness, the dark black color of Kali symbolizes the color from which everything was born. Her right hand side arms she shows the Abhaya mudra(gesture of fearlessness) and Vara mudra (gesture of welcome and charity) respectively . But on the other arm in left side she holds a bloody scimitar and a severed head depicting destruction and end of ego.

Kali as the Symbol of Mother Nature

 

The name Kali means Kala or force of time. When there were neither the creation, nor the sun, the moon, the planets, and the earth, there was only darkness and everything was created from the darkness. The Dark appearance of kali represents the darkness from which everything was born. Her complexion is deep blue, like the sky and ocean water as blue. As she is also the goddess of Preservation Kali is worshiped as mother to preserve the nature.Kali is standing calm on Shiva, her appearance represents the preservation of mother nature. Her free, long and black hair represents nature's freedom from civilization. Under the third eye of kali, the signs of both sun, moon and fire are visible which represent the driving forces of nature.

 

SHIVA IN KALI ICONOGRAPHY

In both these images she is shown standing on the prone, inert or dead body of Shiva. There is a legend for the reason behind her standing on what appears to be Shiva's corpse, which translates as follows:

 

Once Kali had destroyed all the demons in battle, she began a terrific dance out of the sheer joy of victory. All the worlds or lokas began to tremble and sway under the impact of her dance. So, at the request of all the Gods, Shiva himself asked her to desist from this behavior. However, she was too intoxicated to listen. Hence, Shiva lay like a corpse among the slain demons in order to absorb the shock of the dance into himself. When Kali eventually stepped upon Shiva, she realized she was trampling and hurting her husband and bit her tongue in shame.

 

The story described here is a popular folk tale and not described or hinted in any of the puranas. The puranic interpretation is as follows:

 

Once, Parvati asks Shiva to chose the one form among her 10 forms which he likes most. To her surprise, Shiva reveals that he is most comfortable with her Kali form, in which she is bereft of her jewellery, her human-form, her clothes, her emotions and where she is only raw, chaotic energy, where she is as terrible as time itself and even greater than time. As Parvati takes the form of Kali, Shiva lies at her feet and requests her to place her foot on his chest, upon his heart. Once in this form, Shiva requests her to have this place, below her feet in her iconic image which would be worshiped throughout.

 

This idea has been explored in the Devi-Bhagavata Purana [28] and is most popular in the Shyama Sangeet, devotional songs to Kali from the 12th to 15th centuries.

 

The Tantric interpretation of Kali standing on top of her husband is as follows:

 

The Shiv tattava (Divine Consciousness as Shiva) is inactive, while the Shakti tattava (Divine Energy as Kali) is active. Shiva and Kali represent Brahman, the Absolute pure consciousness which is beyond all names, forms and activities. Kali, on the other hand, represents the potential (and manifested) energy responsible for all names, forms and activities. She is his Shakti, or creative power, and is seen as the substance behind the entire content of all consciousness. She can never exist apart from Shiva or act independently of him, just as Shiva remains a mere corpse without Kali i.e., Shakti, all the matter/energy of the universe, is not distinct from Shiva, or Brahman, but is rather the dynamic power of Brahman. Hence, Kali is Para Brahman in the feminine and dynamic aspect while Shiva is the male aspect and static. She stands as the absolute basis for all life, energy and beneath her feet lies, Shiva, a metaphor for mass, which cannot retain its form without energy.

 

While this is an advanced concept in monistic Shaktism, it also agrees with the Nondual Trika philosophy of Kashmir, popularly known as Kashmir Shaivism and associated most famously with Abhinavagupta. There is a colloquial saying that "Shiva without Shakti is Shava" which means that without the power of action (Shakti) that is Mahakali (represented as the short "i" in Devanagari) Shiva (or consciousness itself) is inactive; Shava means corpse in Sanskrit and the play on words is that all Sanskrit consonants are assumed to be followed by a short letter "a" unless otherwise noted. The short letter "i" represents the female power or Shakti that activates Creation. This is often the explanation for why She is standing on Shiva, who is either Her husband and complement in Shaktism or the Supreme Godhead in Shaivism.

 

To properly understand this complex Tantric symbolism it is important to remember that the meaning behind Shiva and Kali does not stray from the non-dualistic parlance of Shankara or the Upanisads. According to both the Mahanirvana and Kularnava Tantras, there are two distinct ways of perceiving the same absolute reality. The first is a transcendental plane which is often described as static, yet infinite. It is here that there is no matter, there is no universe and only consciousness exists. This form of reality is known as Shiva, the absolute Sat-Chit-Ananda - existence, knowledge and bliss. The second is an active plane, an immanent plane, the plane of matter, of Maya, i.e., where the illusion of space-time and the appearance of an actual universe does exist. This form of reality is known as Kali or Shakti, and (in its entirety) is still specified as the same Absolute Sat-Chit-Ananda. It is here in this second plane that the universe (as we commonly know it) is experienced and is described by the Tantric seer as the play of Shakti, or God as Mother Kali.

 

From a Tantric perspective, when one meditates on reality at rest, as absolute pure consciousness (without the activities of creation, preservation or dissolution) one refers to this as Shiva or Brahman. When one meditates on reality as dynamic and creative, as the Absolute content of pure consciousness (with all the activities of creation, preservation or dissolution) one refers to it as Kali or Shakti. However, in either case the yogini or yogi is interested in one and the same reality - the only difference being in name and fluctuating aspects of appearance. It is this which is generally accepted as the meaning of Kali standing on the chest of Shiva.

 

Although there is often controversy surrounding the images of divine copulation, the general consensus is benign and free from any carnal impurities in its substance. In Tantra the human body is a symbol for the microcosm of the universe; therefore sexual process is responsible for the creation of the world. Although theoretically Shiva and Kali (or Shakti) are inseparable, like fire and its power to burn, in the case of creation they are often seen as having separate roles. With Shiva as male and Kali as female it is only by their union that creation may transpire. This reminds us of the prakrti and purusa doctrine of Samkhya wherein prakāśa- vimarśa has no practical value, just as without prakrti, purusa is quite inactive. This (once again) stresses the interdependencies of Shiva and Shakti and the vitality of their union.

 

Gopi Krishna proposed that Kali standing on the dead Shiva or Shava (Sanskrit for dead body) symbolised the helplessness of a person undergoing the changing process (psychologically and physiologically) in the body conducted by the Kundalini Shakti.

 

DEVELOPMENT

In the later traditions, Kali has become inextricably linked with Shiva. The unleashed form of Kali often becomes wild and uncontrollable, and only Shiva is able to tame her just as only Kali can tame Shiva. This is both because she is often a transformed version of one of his consorts and because he is able to match her wildness.

 

The ancient text of Kali Kautuvam describes her competition with Shiva in dance, from which the sacred 108 Karanas appeared. Shiva won the competition by acting the urdva tandava, one of the Karanas, by raising his feet to his head. Other texts describe Shiva appearing as a crying infant and appealing to her maternal instincts. While Shiva is said to be able to tame her, the iconography often presents her dancing on his fallen body, and there are accounts of the two of them dancing together, and driving each other to such wildness that the world comes close to unravelling.

 

Shiva's involvement with Tantra and Kali's dark nature have led to her becoming an important Tantric figure. To the Tantric worshippers, it was essential to face her Curse, the terror of death, as willingly as they accepted Blessings from her beautiful, nurturing, maternal aspect. For them, wisdom meant learning that no coin has only one side: as death cannot exist without life, so life cannot exist without death. Kali's role sometimes grew beyond that of a chaos - which could be confronted - to that of one who could bring wisdom, and she is given great metaphysical significance by some Tantric texts. The Nirvāna-tantra clearly presents her uncontrolled nature as the Ultimate Reality, claiming that the trimurti of Brahma, Vishnu and Rudra arise and disappear from her like bubbles from the sea. Although this is an extreme case, the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra declare her the svarupa (own-being) of the Mahadevi (the great Goddess, who is in this case seen as the combination of all devis).The final stage of development is the worshipping of Kali as the Great Mother, devoid of her usual violence. This practice is a break from the more traditional depictions. The pioneers of this tradition are the 18th century Shakta poets such as Ramprasad Sen, who show an awareness of Kali's ambivalent nature. Ramakrishna, the 19th century Bengali saint, was also a great devotee of Kali; the western popularity of whom may have contributed to the more modern, equivocal interpretations of this Goddess. Rachel McDermott's work, however, suggests that for the common, modern worshipper, Kali is not seen as fearful, and only those educated in old traditions see her as having a wrathful component. Some credit to the development of Devi must also be given to Samkhya. Commonly referred to as the Devi of delusion, Mahamaya or Durga, acting in the confines of (but not being bound by) the nature of the three gunas, takes three forms: Maha-Kali, Maha-Lakshmi and Maha-Saraswati, being her tamas-ika, rajas-ika and sattva-ika forms. In this sense, Kali is simply part of a larger whole.

 

Like Sir John Woodroffe and Georg Feuerstein, many Tantric scholars (as well as sincere practitioners) agree that, no matter how propitious or appalling you describe them, Shiva and Devi are simply recognizable symbols for everyday, abstract (yet tangible) concepts such as perception, knowledge, space-time, causation and the process of liberating oneself from the confines of such things. Shiva, symbolizing pure, absolute consciousness, and Devi, symbolizing the entire content of that consciousness, are ultimately one and the same - totality incarnate, a micro-macro-cosmic amalgamation of all subjects, all objects and all phenomenal relations between the "two." Like man and woman who both share many common, human traits yet at the same time they are still different and, therefore, may also be seen as complementary.

 

Worshippers prescribe various benign and horrific qualities to Devi simply out of practicality. They do this so they may have a variety of symbols to choose from, symbols which they can identify and relate with from the perspective of their own, ever-changing time, place and personal level of unfolding. Just like modern chemists or physicists use a variety of molecular and atomic models to describe what is unperceivable through rudimentary, sensory input, the scientists of ontology and epistemology must do the same. One of the underlying distinctions of Tantra, in comparison to other religions, is that it allows the devotee the liberty to choose from a vast array of complementary symbols and rhetoric which suit one's evolving needs and tastes. From an aesthetic standpoint, nothing is interdict and nothing is orthodox. In this sense, the projection of some of Devi's more gentle qualities onto Kali is not sacrilege and the development of Kali really lies in the practitioner, not the murthi.

 

A TIME magazine article of October 27, 1947, used Kali as a symbol and metaphor for the human suffering in British India during its partition that year. In 1971, Ms. Magazine used an image of Kali, her multiple arms juggling modern tasks, as a symbol of modern womanhood on its inaugural issue.

 

Swami Vivekananda wrote his favorite poem Kali the Mother in 1898.

 

KALI IN NEOPAGAN AND NEW AGE PRACTICE

An academic study of Western Kali enthusiasts noted that, "as shown in the histories of all cross-cultural religious transplants, Kali devotionalism in the West must take on its own indigenous forms if it is to adapt to its new environment."[60] The adoption of Kali by the West has raised accusations of cultural appropriation:

 

A variety of writers and thinkers have found Kali an exciting figure for reflection and exploration, notably feminists and participants in New Age spirituality who are attracted to goddess worship. [For them], Kali is a symbol of wholeness and healing, associated especially with repressed female power and sexuality. [However, such interpretations often exhibit] confusion and misrepresentation, stemming from a lack of knowledge of Hindu history among these authors, [who only rarely] draw upon materials written by scholars of the Hindu religious tradition. The majority instead rely chiefly on other popular feminist sources, almost none of which base their interpretations on a close reading of Kali's Indian background. The most important issue arising from this discussion - even more important than the question of 'correct' interpretation - concerns the adoption of other people's religious symbols. It is hard to import the worship of a goddess from another culture: religious associations and connotations have to be learned, imagined or intuited when the deep symbolic meanings embedded in the native culture are not available.

 

INCARNATIONS OF KALI

Draupadi, Wife of Pandavas, was an avatar of Kali, who born to assist Lord Krishna to destroy arrogant kings of India. There is a temple dedicated to this incarnation at Banni Mata Temple at Himachal Pradesh. The vedic deity Nirriti or the Puranic deity Alakshmi is often considered as incarnations of Kali.

 

WIKIPEDIA

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

I may not have posted this pic at all if not for my dear friend Phyllis. She just posted a pic of a honeybee on a flower, a simple pic but with a true insite to there demise.Read the poster to my left. It's so true!

NASA image acquired June 24, 2007

 

Part of the U.S. Virgin Islands, the island of St. John incorporates both a land-based national park and a sea-based national monument. First established in 1956 and later expanded by the U.S. Congress, the Virgin Islands National Park covers more than 7,000 acres (2,800 hectares). Coral Reef National Monument—established ten years ago this week by presidential proclamation—encompasses submerged lands within 3 miles (5 kilometers) of the St. John coast.

 

The Advanced Land Imager (ALI) on NASA’s Earth Observing-1 (EO-1) satellite captured this natural-color image of the eastern half of St. John on June 24, 2007. Settlements—marked by curving roads and light roofs—line the irregular coastline. The pinpoints of white in Coral Bay are probably boats.

 

The island’s interior is carpeted with varying shades of green. Temperatures vary little throughout the year, ranging between the low 80s to mid-90s Fahrenheit (mid-20s to mid-30s Celsius). Although the vegetation appears lush, much of it is second-generation growth, according to the U.S. National Park Service (NPS). During the colonial period, large swaths of the island were clear-cut to accommodate sugar cane production.

 

Along the fringes of the island, the complex seafloor of varying depths colors the water different shades of blue. The Coral Reef National Monument is comprised of interdependent and fragile ecosystems, including coral reefs, mangrove forests, and beds of seagrass. In addition to these ecosystems, the monument incorporates historical artifacts such as shipwrecks, which are rare in the Caribbean.

 

NASA Earth Observatory image created by Jesse Allen and Robert Simmon, using EO-1 ALI data provided courtesy of the NASA EO-1 team. Caption by Michon Scott.

 

Instrument: EO-1 - ALI

 

Credit: NASA Earth Observatory

 

Go here to download the full high res file: earthobservatory.nasa.gov/images/imagerecords/48000/48615...

 

NASA Goddard Space Flight Center enables NASA’s mission through four scientific endeavors: Earth Science, Heliophysics, Solar System Exploration, and Astrophysics. Goddard plays a leading role in NASA’s accomplishments by contributing compelling scientific knowledge to advance the Agency’s mission.

 

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Have a great Sunday.......enjoy the Super Bowl for those of you who are football fans or at least enjoy the commercials :-) As always, thanks for your wonderful comments & support. View On Black

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Kālī, also known as Kālikā (Sanskrit: कालिका), is the Hindu goddess associated with empowerment, shakti. She is the fierce aspect of the goddess Durga (Parvati). The name Kali comes from kāla, which means black, time, death, lord of death: Shiva. Since Shiva is called Kāla— the eternal time — the name of Kālī, his consort, also means "Time" or "Death" (as in "time has come"). Hence, Kāli is the Goddess of Time and Change. Although sometimes presented as dark and violent, her earliest incarnation as a figure of annihilation of evil forces still has some influence. Various Shakta Hindu cosmologies, as well as Shākta Tantric beliefs, worship her as the ultimate reality or Brahman. Comparatively recent devotional movements largely conceive Kāli as a benevolent mother goddess. Kālī is represented as the consort of Lord Shiva, on whose body she is often seen standing. Shiva lies in the path of Kali, whose foot on Shiva subdues her anger.

 

ETYMOLOGY

Kālī is the feminine form of kālam ("black, dark coloured"). Kāla primarily means "time" but also means "black" in honor of being the first creation before light itself. Kālī means "the black one" and refers to her being the entity of "time" or "beyond time." Kāli is strongly associated with Shiva, and Shaivas derive the masculine Kāla (an epithet of Shiva) to come from her feminine name. A nineteenth-century Sanskrit dictionary, the Shabdakalpadrum, states: कालः शिवः। तस्य पत्नीति - काली। kālaḥ śivaḥ। tasya patnīti kālī - "Shiva is Kāla, thus, his consort is Kāli" referring to Devi Parvathi being a manifestation of Devi MahaKali.

 

Other names include Kālarātri ("black night"), as described above, and Kālikā ("relating to time"). Coburn notes that the name Kālī can be used as a proper name, or as a description of color.

 

Kāli's association with darkness stands in contrast to her consort, Shiva, who manifested after her in creation, and who symbolises the rest of creation after Time is created. In his supreme awareness of Maya, his body is covered by the white ashes of the cremation ground (Sanskrit: śmaśāna) where he meditates, and with which Kāli is also associated, as śmaśāna-kālī.

 

ORIGINS

Hugh Urban notes that although the word Kālī appears as early as the Atharva Veda, the first use of it as a proper name is in the Kathaka Grhya Sutra (19.7). Kali is the name of one of the seven tongues of Agni, the [Rigvedic] God of Fire, in the Mundaka Upanishad (2:4), but it is unlikely that this refers to the goddess. The first appearance of Kāli in her present form is in the Sauptika Parvan of the Mahabharata (10.8.64). She is called Kālarātri (literally, "black night") and appears to the Pandava soldiers in dreams, until finally she appears amidst the fighting during an attack by Drona's son Ashwatthama. She most famously appears in the sixth century Devi Mahatmyam as one of the shaktis of Mahadevi, and defeats the demon Raktabija ("Bloodseed"). The tenth-century Kalika Purana venerates Kāli as the ultimate reality.

 

According to David Kinsley, Kāli is first mentioned in Hinduism as a distinct goddess around 600 CE, and these texts "usually place her on the periphery of Hindu society or on the battlefield." She is often regarded as the Shakti of Shiva, and is closely associated with him in various Puranas. The Kalika Purana depicts her as the "Adi Shakti" (Fundamental Power) and "Para Prakriti" or beyond nature.

 

WORSHIP & MANTRA

Kali could be considered a general concept, like Durga, and is mostly worshiped in the Kali Kula sect of worship. The closest way of direct worship is Maha Kali or Bhadra Kali (Bhadra in Sanskrit means 'gentle'). Kali is worshiped as one of the 10 Mahavidya forms of Adi Parashakti (Goddess Durga) or Bhagavathy according to the region. The mantra for worship is called Devi Argala Stotram.

Sanskrit: सर्वमङ्गलमाङ्गल्ये शिवे सर्वार्थसाधिके । शरण्ये त्र्यम्बके गौरि नारायणि नमोऽस्तु ते ॥

 

ॐ जयंती मंगल काली भद्रकाली कपालिनी । दुर्गा क्षमा शिवा धात्री स्वाहा स्वधा नमोऽस्तु‍ते ॥

(Sarvamaṅgalamāṅgalyē śivē sarvārthasādhikē . śaraṇyē tryambakē gauri nārāyaṇi namō'stu tē.

Oṃ jayantī mangala kālī bhadrakālī kapālinī . durgā kṣamā śivā dhātrī svāhā svadhā namō'stu‍tē.)

 

TANTRA

Goddesses play an important role in the study and practice of Tantra Yoga, and are affirmed to be as central to discerning the nature of reality as are the male deities. Although Parvati is often said to be the recipient and student of Shiva's wisdom in the form of Tantras, it is Kāli who seems to dominate much of the Tantric iconography, texts, and rituals. In many sources Kāli is praised as the highest reality or greatest of all deities. The Nirvana-tantra says the gods Brahma, Vishnu, and Shiva all arise from her like bubbles in the sea, ceaselessly arising and passing away, leaving their original source unchanged. The Niruttara-tantra and the Picchila-tantra declare all of Kāli's mantras to be the greatest and the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra all proclaim Kāli vidyas (manifestations of Mahadevi, or "divinity itself"). They declare her to be an essence of her own form (svarupa) of the Mahadevi.

 

In the Mahanirvana-tantra, Kāli is one of the epithets for the primordial sakti, and in one passage Shiva praises her:

 

At the dissolution of things, it is Kāla [Time] Who will devour all, and by reason of this He is called Mahākāla [an epithet of Lord Shiva], and since Thou devourest Mahākāla Himself, it is Thou who art the Supreme Primordial Kālika. Because Thou devourest Kāla, Thou art Kāli, the original form of all things, and because Thou art the Origin of and devourest all things Thou art called the Adya [the Primordial One]. Re-assuming after Dissolution Thine own form, dark and formless, Thou alone remainest as One ineffable and inconceivable. Though having a form, yet art Thou formless; though Thyself without beginning, multiform by the power of Maya, Thou art the Beginning of all, Creatrix, Protectress, and Destructress that Thou art.

 

The figure of Kāli conveys death, destruction, and the consuming aspects of reality. As such, she is also a "forbidden thing", or even death itself. In the Pancatattva ritual, the sadhaka boldly seeks to confront Kali, and thereby assimilates and transforms her into a vehicle of salvation. This is clear in the work of the Karpuradi-stotra, a short praise of Kāli describing the Pancatattva ritual unto her, performed on cremation grounds. (Samahana-sadhana)

 

He, O Mahākāli who in the cremation-ground, naked, and with dishevelled hair, intently meditates upon Thee and recites Thy mantra, and with each recitation makes offering to Thee of a thousand Akanda flowers with seed, becomes without any effort a Lord of the earth. Oh Kāli, whoever on Tuesday at midnight, having uttered Thy mantra, makes offering even but once with devotion to Thee of a hair of his Shakti [his energy/female companion] in the cremation-ground, becomes a great poet, a Lord of the earth, and ever goes mounted upon an elephant.

 

The Karpuradi-stotra clearly indicates that Kāli is more than a terrible, vicious, slayer of demons who serves Durga or Shiva. Here, she is identified as the supreme mistress of the universe, associated with the five elements. In union with Lord Shiva, she creates and destroys worlds. Her appearance also takes a different turn, befitting her role as ruler of the world and object of meditation. In contrast to her terrible aspects, she takes on hints of a more benign dimension. She is described as young and beautiful, has a gentle smile, and makes gestures with her two right hands to dispel any fear and offer boons. The more positive features exposed offer the distillation of divine wrath into a goddess of salvation, who rids the sadhaka of fear. Here, Kali appears as a symbol of triumph over death.

 

BENGALI TRADITION

Kali is also a central figure in late medieval Bengali devotional literature, with such devotees as Ramprasad Sen (1718–75). With the exception of being associated with Parvati as Shiva's consort, Kāli is rarely pictured in Hindu legends and iconography as a motherly figure until Bengali devotions beginning in the early eighteenth century. Even in Bengāli tradition her appearance and habits change little, if at all.

 

The Tantric approach to Kāli is to display courage by confronting her on cremation grounds in the dead of night, despite her terrible appearance. In contrast, the Bengali devotee appropriates Kāli's teachings adopting the attitude of a child, coming to love her unreservedly. In both cases, the goal of the devotee is to become reconciled with death and to learn acceptance of the way that things are. These themes are well addressed in Rāmprasād's work. Rāmprasād comments in many of his other songs that Kāli is indifferent to his wellbeing, causes him to suffer, brings his worldly desires to nothing and his worldly goods to ruin. He also states that she does not behave like a mother should and that she ignores his pleas:

 

Can mercy be found in the heart of her who was born of the stone? [a reference to Kali as the daughter of Himalaya]

Were she not merciless, would she kick the breast of her lord?

Men call you merciful, but there is no trace of mercy in you, Mother.

You have cut off the heads of the children of others, and these you wear as a garland around your neck.

It matters not how much I call you "Mother, Mother." You hear me, but you will not listen.

 

To be a child of Kāli, Rāmprasād asserts, is to be denied of earthly delights and pleasures. Kāli is said to refrain from giving that which is expected. To the devotee, it is perhaps her very refusal to do so that enables her devotees to reflect on dimensions of themselves and of reality that go beyond the material world.

 

A significant portion of Bengali devotional music features Kāli as its central theme and is known as Shyama Sangeet ("Music of the Night"). Mostly sung by male vocalists, today even women have taken to this form of music. One of the finest singers of Shyāma Sāngeet is Pannalal Bhattacharya.

 

In Bengal, Kāli is venerated in the festival Kali Puja, the new moon day of Ashwin month which coincides with Diwali festival.

 

In a unique form of Kāli worship, Shantipur worships Kāli in the form of a hand painted image of the deity known as Poteshwari (meaning the deity drawn on a piece of cloth).

 

LEGENDS

SLAYER OF RAKTABIJA

In Kāli's most famous legend, Devi Durga (Adi Parashakti) and her assistants, the Matrikas, wound the demon Raktabija, in various ways and with a variety of weapons in an attempt to destroy him. They soon find that they have worsened the situation for with every drop of blood that is dripped from Raktabija he reproduces a clone of himself. The battlefield becomes increasingly filled with his duplicates. Durga, in need of help, summons Kāli to combat the demons. It is said, in some versions, that Goddess Durga actually assumes the form of Goddess Kāli at this time. The Devi Mahatmyam describes:

 

Out of the surface of her (Durga's) forehead, fierce with frown, issued suddenly Kali of terrible countenance, armed with a sword and noose. Bearing the strange khatvanga (skull-topped staff ), decorated with a garland of skulls, clad in a tiger's skin, very appalling owing to her emaciated flesh, with gaping mouth, fearful with her tongue lolling out, having deep reddish eyes, filling the regions of the sky with her roars, falling upon impetuously and slaughtering the great asuras in that army, she devoured those hordes of the foes of the devas.

 

Kali destroys Raktabija by sucking the blood from his body and putting the many Raktabija duplicates in her gaping mouth. Pleased with her victory, Kali then dances on the field of battle, stepping on the corpses of the slain. In the Devi Mahatmya version of this story, Kali is also described as a Matrika and as a Shakti or power of Devi. She is given the epithet Cāṃuṇḍā (Chamunda), i.e. the slayer of the demons Chanda and Munda. Chamunda is very often identified with Kali and is very much like her in appearance and habit.

 

DAKSHINA KALI

In her most famous pose as Daksinakali, popular legends say that Kali, becoming drunk on the blood of her victims on the battlefield, dances with destructive frenzy. She is about to destroy the whole universe when, urged by all the gods, Shiva lies in her way to stop her. In her fury, she fails to see the body of Shiva lying amongst the corpses on the battlefield and steps upon his chest. Realizing Shiva lies beneath her feet, her anger is pacified and she calms her fury. Though not included in any of the puranas, popular legends state that Kali was ashamed at the prospect of keeping her husband beneath her feet and thus stuck her tongue out in shame. The Devi-Bhagavata Purana, which goes into great depths about the goddess Kali, reveals the tongue's actual symbolism.

 

The characteristic icons that depict Kali are the following; unbridled matted hair, open blood shot eyes, open mouth and a drooping tongue; in her hands, she holds a Khadga (bent sword or scimitar) and a human head; she has a girdle of human hands across her waist and an enchanted Shiva lies beneath her feet. Each of these icons represent a deep philosophical epithet. The drooping out-stuck tongue represents her blood-thirst. Lord Shiva beneath her feet represents matter, as Kali is undoubtedly the primeval energy. The depiction of Kali on Shiva shows that without energy, matter lies "dead". This concept has been simplified to a folk-tale depicting a wife placing her foot on her husband and sticking her tongue out in shame. In tantric contexts, the tongue is seen to denote the element (guna) of rajas (energy and action) controlled by sattva.

 

If Kali steps on Shiva with her right foot and holds the sword in her left hand, she is considered to be Dakshina Kali. The Dakshina Kali Temple has important religious associations with the Jagannath Temple and it is believed that Daksinakali is the guardian of the kitchen of the Lord Jagannath Temple. Puranic tradition says that in Puri, Lord Jagannath is regarded as Daksinakalika. Goddess Dakshinakali plays an important role in the 'Niti' of Saptapuri Amavasya.

 

One South Indian tradition tells of a dance contest between Shiva and Kali. After defeating the two demons Sumbha and Nisumbha, Kali takes up residence in the forest of Thiruvalankadu or Thiruvalangadu. She terrorizes the surrounding area with her fierce, disruptive nature. One of Shiva's devotees becomes distracted while performing austerities, and asks Shiva to rid the forest of the destructive goddess. When Shiva arrives, Kali threatens him, claiming the territory as her own. Shiva challenges Kali to a dance contest; both of them dance and Kali matches Shiva in every step that he takes until Shiva takes the "Urdhvatandava" step, by vertically raising his right leg. Kali refuses to perform this step, which would not befit her as a woman, and became pacified.

 

SMASHAN KALI

If the Kali steps out with the left foot and holds the sword in her right hand, she is the terrible form of Mother, the Smashan Kali of the cremation ground. She is worshiped by tantrics, the followers of Tantra, who believe that one's spiritual discipline practiced in a smashan (cremation ground) brings success quickly. Sarda Devi, the consort of Ramakrishna Paramhansa, worshipped Smashan Kali at Dakshineshwar.

 

MATERNAL KALI

Another legend depicts the infant Shiva calming Kali. In this similar story, Kali has defeated her enemies on the battlefield and begun to dance out of control, drunk on the blood of the slain. To calm her down and to protect the stability of the world, Shiva is sent to the battlefield, as an infant, crying aloud. Seeing the child's distress, Kali ceases dancing to care for the helpless infant. She picks him up, kisses his head, and proceeds to breast feed the infant Shiva. This legend is notable because it shows Kali in her benevolent, maternal aspect, with which she is not usually identified.

 

MAHAKALI

Mahakali (Sanskrit: Mahākālī, Devanagari: महाकाली), literally translated as Great Kali, is sometimes considered as a greater form of Kali, identified with the Ultimate reality of Brahman. It can also be used as an honorific of the Goddess Kali, signifying her greatness by the prefix "Mahā-". Mahakali, in Sanskrit, is etymologically the feminized variant of Mahakala or Great Time (which is interpreted also as Death), an epithet of the God Shiva in Hinduism. Mahakali is the presiding Goddess of the first episode of the Devi Mahatmya. Here she is depicted as Devi in her universal form as Shakti. Here Devi serves as the agent who allows the cosmic order to be restored.

 

Kali is depicted in the Mahakali form as having ten heads, ten arms, and ten legs. Each of her ten hands is carrying a various implement which vary in different accounts, but each of these represent the power of one of the Devas or Hindu Gods and are often the identifying weapon or ritual item of a given Deva. The implication is that Mahakali subsumes and is responsible for the powers that these deities possess and this is in line with the interpretation that Mahakali is identical with Brahman. While not displaying ten heads, an "ekamukhi" or one headed image may be displayed with ten arms, signifying the same concept: the powers of the various Gods come only through Her grace.

 

ICONOGRAPHY

Kali is portrayed mostly in two forms: the popular four-armed form and the ten-armed Mahakali form. In both of her forms, she is described as being black in color but is most often depicted as blue in popular Indian art. Her eyes are described as red with intoxication, and in absolute rage, her hair is shown disheveled, small fangs sometimes protrude out of her mouth, and her tongue is lolling. She is often shown naked or just wearing a skirt made of human arms and a garland of human heads. She is also accompanied by serpents and a jackal while standing on a seemingly dead Shiva, usually right foot forward to symbolize the more popular Dakshinamarga or right-handed path, as opposed to the more infamous and transgressive Vamamarga or left-handed path.

 

In the ten-armed form of Mahakali she is depicted as shining like a blue stone. She has ten faces and ten feet and three eyes. She has ornaments decked on all her limbs. There is no association with Shiva.

 

The Kalika Purana describes Kali as possessing a soothing dark complexion, as perfectly beautiful, riding a lion, four-armed, holding a sword and blue lotuses, her hair unrestrained, body firm and youthful.

 

In spite of her seemingly terrible form, Kali Ma is often considered the kindest and most loving of all the Hindu goddesses, as she is regarded by her devotees as the Mother of the whole Universe. And because of her terrible form, she is also often seen as a great protector. When the Bengali saint Ramakrishna once asked a devotee why one would prefer to worship Mother over him, this devotee rhetorically replied, "Maharaj, when they are in trouble your devotees come running to you. But, where do you run when you are in trouble?"

 

According to Ramakrishna, darkness is the Ultimate Mother, or Kali:

 

My Mother is the principle of consciousness. She is Akhanda Satchidananda; indivisible Reality, Awareness, and Bliss. The night sky between the stars is perfectly black. The waters of the ocean depths are the same; The infinite is always mysteriously dark. This inebriating darkness is my beloved Kali.

 

SRI RAMAKRISHNA

This is clear in the works of such contemporary artists as Charles Wish, and Tyeb Mehta, who sometimes take great liberties with the traditional, accepted symbolism, but still demonstrate a true reverence for the Shakta sect.

 

POPULAR FORM

Classic depictions of Kali share several features, as follows:

 

Kali's most common four armed iconographic image shows each hand carrying variously a sword, a trishul (trident), a severed head and a bowl or skull-cup (kapala) catching the blood of the severed head.

 

Two of these hands (usually the left) are holding a sword and a severed head. The Sword signifies Divine Knowledge and the Human Head signifies human Ego which must be slain by Divine Knowledge in order to attain Moksha. The other two hands (usually the right) are in the abhaya (fearlessness) and varada (blessing) mudras, which means her initiated devotees (or anyone worshipping her with a true heart) will be saved as she will guide them here and in the hereafter.

 

She has a garland consisting of human heads, variously enumerated at 108 (an auspicious number in Hinduism and the number of countable beads on a Japa Mala or rosary for repetition of Mantras) or 51, which represents Varnamala or the Garland of letters of the Sanskrit alphabet, Devanagari. Hindus believe Sanskrit is a language of dynamism, and each of these letters represents a form of energy, or a form of Kali. Therefore she is generally seen as the mother of language, and all mantras.

 

She is often depicted naked which symbolizes her being beyond the covering of Maya since she is pure (nirguna) being-consciousness-bliss and far above prakriti. She is shown as very dark as she is brahman in its supreme unmanifest state. She has no permanent qualities - she will continue to exist even when the universe ends. It is therefore believed that the concepts of color, light, good, bad do not apply to her - she is the pure, un-manifested energy, the Adi-shakti.

 

SHIVA IN KALI ICONOGRAPHY

In both these images she is shown standing on the prone, inert or dead body of Shiva. There is a legend for the reason behind her standing on what appears to be Shiva's corpse, which translates as follows:

 

Once Kali had destroyed all the demons in battle, she began a terrific dance out of the sheer joy of victory. All the worlds or lokas began to tremble and sway under the impact of her dance. So, at the request of all the Gods, Shiva himself asked her to desist from this behavior. However, she was too intoxicated to listen. Hence, Shiva lay like a corpse among the slain demons in order to absorb the shock of the dance into himself. When Kali eventually stepped upon Shiva, she realized she was trampling and hurting her husband and bit her tongue in shame.

 

The story described here is a popular folk tale and not described or hinted in any of the puranas. The puranic interpretation is as follows:

 

Once, Parvati asks Shiva to chose the one form among her 10 forms which he likes most. To her surprise, Shiva reveals that he is most comfortable with her Kali form, in which she is bereft of her jewellery, her human-form, her clothes, her emotions and where she is only raw, chaotic energy, where she is as terrible as time itself and even greater than time. As Parvati takes the form of Kali, Shiva lies at her feet and requests her to place her foot on his chest, upon his heart. Once in this form, Shiva requests her to have this place, below her feet in her iconic image which would be worshiped throughout.

 

This idea has been explored in the Devi-Bhagavata Purana and is most popular in the Shyama Sangeet, devotional songs to Kali from the 12th to 15th centuries.

 

The Tantric interpretation of Kali standing on top of her husband is as follows:

 

The Shiv tattava (Divine Consciousness as Shiva) is inactive, while the Shakti tattava (Divine Energy as Kali) is active. Shiva and Kali represent Brahman, the Absolute pure consciousness which is beyond all names, forms and activities. Kali, on the other hand, represents the potential (and manifested) energy responsible for all names, forms and activities. She is his Shakti, or creative power, and is seen as the substance behind the entire content of all consciousness. She can never exist apart from Shiva or act independently of him, just as Shiva remains a mere corpse without Kali i.e., Shakti, all the matter/energy of the universe, is not distinct from Shiva, or Brahman, but is rather the dynamic power of Brahman. Hence, Kali is Para Brahman in the feminine and dynamic aspect while Shiva is the male aspect and static. She stands as the absolute basis for all life, energy and beneath her feet lies, Shiva, a metaphor for mass, which cannot retain its form without energy.

 

While this is an advanced concept in monistic Shaktism, it also agrees with the Nondual Trika philosophy of Kashmir, popularly known as Kashmir Shaivism and associated most famously with Abhinavagupta. There is a colloquial saying that "Shiva without Shakti is Shava" which means that without the power of action (Shakti) that is Mahakali (represented as the short "i" in Devanagari) Shiva (or consciousness itself) is inactive; Shava means corpse in Sanskrit and the play on words is that all Sanskrit consonants are assumed to be followed by a short letter "a" unless otherwise noted. The short letter "i" represents the female power or Shakti that activates Creation. This is often the explanation for why She is standing on Shiva, who is either Her husband and complement in Shaktism or the Supreme Godhead in Shaivism.

 

To properly understand this complex Tantric symbolism it is important to remember that the meaning behind Shiva and Kali does not stray from the non-dualistic parlance of Shankara or the Upanisads. According to both the Mahanirvana and Kularnava Tantras, there are two distinct ways of perceiving the same absolute reality. The first is a transcendental plane which is often described as static, yet infinite. It is here that there is no matter, there is no universe and only consciousness exists. This form of reality is known as Shiva, the absolute Sat-Chit-Ananda - existence, knowledge and bliss. The second is an active plane, an immanent plane, the plane of matter, of Maya, i.e., where the illusion of space-time and the appearance of an actual universe does exist. This form of reality is known as Kali or Shakti, and (in its entirety) is still specified as the same Absolute Sat-Chit-Ananda. It is here in this second plane that the universe (as we commonly know it) is experienced and is described by the Tantric seer as the play of Shakti, or God as Mother Kali.

 

From a Tantric perspective, when one meditates on reality at rest, as absolute pure consciousness (without the activities of creation, preservation or dissolution) one refers to this as Shiva or Brahman. When one meditates on reality as dynamic and creative, as the Absolute content of pure consciousness (with all the activities of creation, preservation or dissolution) one refers to it as Kali or Shakti. However, in either case the yogini or yogi is interested in one and the same reality - the only difference being in name and fluctuating aspects of appearance. It is this which is generally accepted as the meaning of Kali standing on the chest of Shiva.

 

Although there is often controversy surrounding the images of divine copulation, the general consensus is benign and free from any carnal impurities in its substance. In Tantra the human body is a symbol for the microcosm of the universe; therefore sexual process is responsible for the creation of the world. Although theoretically Shiva and Kali (or Shakti) are inseparable, like fire and its power to burn, in the case of creation they are often seen as having separate roles. With Shiva as male and Kali as female it is only by their union that creation may transpire. This reminds us of the prakrti and purusa doctrine of Samkhya wherein prakāśa- vimarśa has no practical value, just as without prakrti, purusa is quite inactive. This (once again) stresses the interdependencies of Shiva and Shakti and the vitality of their union.

 

Gopi Krishna proposed that Kali standing on the dead Shiva or Shava (Sanskrit for dead body) symbolised the helplessness of a person undergoing the changing process (psychologically and physiologically) in the body conducted by the Kundalini Shakti.

 

DEVELOPMENT

In the later traditions, Kali has become inextricably linked with Shiva. The unleashed form of Kali often becomes wild and uncontrollable, and only Shiva is able to tame her just as only Kali can tame Shiva. This is both because she is often a transformed version of one of his consorts and because he is able to match her wildness.

 

The ancient text of Kali Kautuvam describes her competition with Shiva in dance, from which the sacred 108 Karanas appeared. Shiva won the competition by acting the urdva tandava, one of the Karanas, by raising his feet to his head. Other texts describe Shiva appearing as a crying infant and appealing to her maternal instincts. While Shiva is said to be able to tame her, the iconography often presents her dancing on his fallen body, and there are accounts of the two of them dancing together, and driving each other to such wildness that the world comes close to unravelling.

 

Shiva's involvement with Tantra and Kali's dark nature have led to her becoming an important Tantric figure. To the Tantric worshippers, it was essential to face her Curse, the terror of death, as willingly as they accepted Blessings from her beautiful, nurturing, maternal aspect. For them, wisdom meant learning that no coin has only one side: as death cannot exist without life, so life cannot exist without death. Kali's role sometimes grew beyond that of a chaos - which could be confronted - to that of one who could bring wisdom, and she is given great metaphysical significance by some Tantric texts. The Nirvāna-tantra clearly presents her uncontrolled nature as the Ultimate Reality, claiming that the trimurti of Brahma, Vishnu and Rudra arise and disappear from her like bubbles from the sea. Although this is an extreme case, the Yogini-tantra, Kamakhya-tantra and the Niruttara-tantra declare her the svarupa (own-being) of the Mahadevi (the great Goddess, who is in this case seen as the combination of all devis).

 

The final stage of development is the worshipping of Kali as the Great Mother, devoid of her usual violence. This practice is a break from the more traditional depictions. The pioneers of this tradition are the 18th century Shakta poets such as Ramprasad Sen, who show an awareness of Kali's ambivalent nature. Ramakrishna, the 19th century Bengali saint, was also a great devotee of Kali; the western popularity of whom may have contributed to the more modern, equivocal interpretations of this Goddess. Rachel McDermott's work, however, suggests that for the common, modern worshipper, Kali is not seen as fearful, and only those educated in old traditions see her as having a wrathful component. Some credit to the development of Devi must also be given to Samkhya. Commonly referred to as the Devi of delusion, Mahamaya or Durga, acting in the confines of (but not being bound by) the nature of the three gunas, takes three forms: Maha-Kali, Maha-Lakshmi and Maha-Saraswati, being her tamas-ika, rajas-ika and sattva-ika forms. In this sense, Kali is simply part of a larger whole.

 

Like Sir John Woodroffe and Georg Feuerstein, many Tantric scholars (as well as sincere practitioners) agree that, no matter how propitious or appalling you describe them, Shiva and Devi are simply recognizable symbols for everyday, abstract (yet tangible) concepts such as perception, knowledge, space-time, causation and the process of liberating oneself from the confines of such things. Shiva, symbolizing pure, absolute consciousness, and Devi, symbolizing the entire content of that consciousness, are ultimately one and the same - totality incarnate, a micro-macro-cosmic amalgamation of all subjects, all objects and all phenomenal relations between the "two." Like man and woman who both share many common, human traits yet at the same time they are still different and, therefore, may also be seen as complementary.

 

Worshippers prescribe various benign and horrific qualities to Devi simply out of practicality. They do this so they may have a variety of symbols to choose from, symbols which they can identify and relate with from the perspective of their own, ever-changing time, place and personal level of unfolding. Just like modern chemists or physicists use a variety of molecular and atomic models to describe what is unperceivable through rudimentary, sensory input, the scientists of ontology and epistemology must do the same. One of the underlying distinctions of Tantra, in comparison to other religions, is that it allows the devotee the liberty to choose from a vast array of complementary symbols and rhetoric which suit one's evolving needs and tastes. From an aesthetic standpoint, nothing is interdict and nothing is orthodox. In this sense, the projection of some of Devi's more gentle qualities onto Kali is not sacrilege and the development of Kali really lies in the practitioner, not the murthi.

 

A TIME magazine article of October 27, 1947, used Kali as a symbol and metaphor for the human suffering in British India during its partition that year.

 

Swami Vivekananda wrote his favorite poem Kali the Mother in 1898.

 

IN NEW AGE & NEOPAGANISM

An academic study of Western Kali enthusiasts noted that, "as shown in the histories of all cross-cultural religious transplants, Kali devotionalism in the West must take on its own indigenous forms if it is to adapt to its new environment." The adoption of Kali by the West has raised accusations of cultural appropriation:

 

A variety of writers and thinkers have found Kali an exciting figure for reflection and exploration, notably feminists and participants in New Age spirituality who are attracted to goddess worship. Kali is a symbol of wholeness and healing, associated especially with repressed female power and sexuality. [However, such interpretations often exhibit] confusion and misrepresentation, stemming from a lack of knowledge of Hindu history among these authors, draw upon materials written by scholars of the Hindu religious tradition. The majority instead rely chiefly on other popular feminist sources, almost none of which base their interpretations on a close reading of Kali's Indian background. The most important issue arising from this discussion - even more important than the question of 'correct' interpretation - concerns the adoption of other people's religious symbols. It is hard to import the worship of a goddess from another culture: religious associations and connotations have to be learned, imagined or intuited when the deep symbolic meanings embedded in the native culture are not available.

 

WIKIPEDIA

30 Days of Perception - Day 29 - Interdependence

Whenever I prepare a meal or eat a fruit. I think about which country grows that food, how it grows and in which season. How dependent we are on the elements necessary to produce our food like: the quality of the soil, the rain and the sunshine. Then there are the people who harvest our food and the markets that sell the fresh produce. I am certainly grateful for all these different stages and feel a connection to each step which allows me to cook a delicious tasting meal.

Just on one plate, I have pasta and Parmesan cheese from Italy, Chickpeas from Africa or Asia, Greek olives and vegetables from Switzerland and Spain. The basil topping comes from my own homegrown plant.

 

We are all interconnected and interdependent on each other.

The Italian pasta dish is called "Cinque Pi" (Five Ps)

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

The truth shall set you free, or kill you trying to get there.

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Si Vis Pacem Para Bellum

"If You Wish For Peace; Prepare For War"

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“We must hang together, gentlemen...else, we shall most assuredly hang separately.” -- Benjamin Franklin

 

"Those who would give up essential Liberty, to purchase a little temporary Safety, deserve neither Liberty nor Safety." -- Benjamin Franklin

 

"The tree of liberty must be refreshed from time to time with the blood of patriots and tyrants." -- Thomas Jefferson

 

“When the people fear the government, that's tyranny; when the government fears the people, that's freedom.” -- Thomas Jefferson

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Military-Industrial Complex Speech, Dwight D. Eisenhower, 1961

 

Public Papers of the Presidents, Dwight D. Eisenhower, 1960, p. 1035- 1040

 

My fellow Americans:

 

Three days from now, after half a century in the service of our country, I shall lay down the responsibilities of office as, in traditional and solemn ceremony, the authority of the Presidency is vested in my successor.

 

This evening I come to you with a message of leave-taking and farewell, and to share a few final thoughts with you, my countrymen.

 

Like every other citizen, I wish the new President, and all who will labor with him, Godspeed. I pray that the coming years will be blessed with peace and prosperity for all.

 

Our people expect their President and the Congress to find essential agreement on issues of great moment, the wise resolution of which will better shape the future of the Nation.

 

My own relations with the Congress, which began on a remote and tenuous basis when, long ago, a member of the Senate appointed me to West Point, have since ranged to the intimate during the war and immediate post-war period, and, finally, to the mutually interdependent during these past eight years.

 

In this final relationship, the Congress and the Administration have, on most vital issues, cooperated well, to serve the national good rather than mere partisanship, and so have assured that the business of the Nation should go forward. So, my official relationship with the Congress ends in a feeling, on my part, of gratitude that we have been able to do so much together.

 

II.

 

We now stand ten years past the midpoint of a century that has witnessed four major wars among great nations. Three of these involved our own country. Despite these holocausts America is today the strongest, the most influential and most productive nation in the world. Understandably proud of this pre-eminence, we yet realize that America's leadership and prestige depend, not merely upon our unmatched material progress, riches and military strength, but on how we use our power in the interests of world peace and human betterment.

 

III.

 

Throughout America's adventure in free government, our basic purposes have been to keep the peace; to foster progress in human achievement, and to enhance liberty, dignity and integrity among people and among nations. To strive for less would be unworthy of a free and religious people. Any failure traceable to arrogance, or our lack of comprehension or readiness to sacrifice would inflict upon us grievous hurt both at home and abroad.

 

Progress toward these noble goals is persistently threatened by the conflict now engulfing the world. It commands our whole attention, absorbs our very beings. We face a hostile ideology -- global in scope, atheistic in character, ruthless in purpose, and insidious in method. Unhappily the danger is poses promises to be of indefinite duration. To meet it successfully, there is called for, not so much the emotional and transitory sacrifices of crisis, but rather those which enable us to carry forward steadily, surely, and without complaint the burdens of a prolonged and complex struggle -- with liberty the stake. Only thus shall we remain, despite every provocation, on our charted course toward permanent peace and human betterment.

 

Crises there will continue to be. In meeting them, whether foreign or domestic, great or small, there is a recurring temptation to feel that some spectacular and costly action could become the miraculous solution to all current difficulties. A huge increase in newer elements of our defense; development of unrealistic programs to cure every ill in agriculture; a dramatic expansion in basic and applied research -- these and many other possibilities, each possibly promising in itself, may be suggested as the only way to the road we wish to travel.

 

But each proposal must be weighed in the light of a broader consideration: the need to maintain balance in and among national programs -- balance between the private and the public economy, balance between cost and hoped for advantage -- balance between the clearly necessary and the comfortably desirable; balance between our essential requirements as a nation and the duties imposed by the nation upon the individual; balance between actions of the moment and the national welfare of the future. Good judgment seeks balance and progress; lack of it eventually finds imbalance and frustration.

 

The record of many decades stands as proof that our people and their government have, in the main, understood these truths and have responded to them well, in the face of stress and threat. But threats, new in kind or degree, constantly arise. I mention two only.

 

IV.

 

A vital element in keeping the peace is our military establishment. Our arms must be mighty, ready for instant action, so that no potential aggressor may be tempted to risk his own destruction.

 

Our military organization today bears little relation to that known by any of my predecessors in peacetime, or indeed by the fighting men of World War II or Korea.

 

Until the latest of our world conflicts, the United States had no armaments industry. American makers of plowshares could, with time and as required, make swords as well. But now we can no longer risk emergency improvisation of national defense; we have been compelled to create a permanent armaments industry of vast proportions. Added to this, three and a half million men and women are directly engaged in the defense establishment. We annually spend on military security more than the net income of all United States corporations.

 

This conjunction of an immense military establishment and a large arms industry is new in the American experience. The total influence -- economic, political, even spiritual -- is felt in every city, every State house, every office of the Federal government. We recognize the imperative need for this development. Yet we must not fail to comprehend its grave implications. Our toil, resources and livelihood are all involved; so is the very structure of our society.

 

In the councils of government, we must guard against the acquisition of unwarranted influence, whether sought or unsought, by the military industrial complex. The potential for the disastrous rise of misplaced power exists and will persist.

 

We must never let the weight of this combination endanger our liberties or democratic processes. We should take nothing for granted. Only an alert and knowledgeable citizenry can compel the proper meshing of the huge industrial and military machinery of defense with our peaceful methods and goals, so that security and liberty may prosper together.

 

Akin to, and largely responsible for the sweeping changes in our industrial-military posture, has been the technological revolution during recent decades.

 

In this revolution, research has become central; it also becomes more formalized, complex, and costly. A steadily increasing share is conducted for, by, or at the direction of, the Federal government.

 

Today, the solitary inventor, tinkering in his shop, has been overshadowed by task forces of scientists in laboratories and testing fields. In the same fashion, the free university, historically the fountainhead of free ideas and scientific discovery, has experienced a revolution in the conduct of research. Partly because of the huge costs involved, a government contract becomes virtually a substitute for intellectual curiosity. For every old blackboard there are now hundreds of new electronic computers.

 

The prospect of domination of the nation's scholars by Federal employment, project allocations, and the power of money is ever present

 

and is gravely to be regarded.

 

Yet, in holding scientific research and discovery in respect, as we should, we must also be alert to the equal and opposite danger that public policy could itself become the captive of a scientific technological elite.

 

It is the task of statesmanship to mold, to balance, and to integrate these and other forces, new and old, within the principles of our democratic system -- ever aiming toward the supreme goals of our free society.

 

V.

 

Another factor in maintaining balance involves the element of time. As we peer into society's future, we -- you and I, and our government -- must avoid the impulse to live only for today, plundering, for our own ease and convenience, the precious resources of tomorrow. We cannot mortgage the material assets of our grandchildren without risking the loss also of their political and spiritual heritage. We want democracy to survive for all generations to come, not to become the insolvent phantom of tomorrow.

 

VI.

 

Down the long lane of the history yet to be written America knows that this world of ours, ever growing smaller, must avoid becoming a community of dreadful fear and hate, and be instead, a proud confederation of mutual trust and respect.

 

Such a confederation must be one of equals. The weakest must come to the conference table with the same confidence as do we, protected as we are by our moral, economic, and military strength. That table, though scarred by many past frustrations, cannot be abandoned for the certain agony of the battlefield.

 

Disarmament, with mutual honor and confidence, is a continuing imperative. Together we must learn how to compose differences, not with arms, but with intellect and decent purpose. Because this need is so sharp and apparent I confess that I lay down my official responsibilities in this field with a definite sense of disappointment. As one who has witnessed the horror and the lingering sadness of war -- as one who knows that another war could utterly destroy this civilization which has been so slowly and painfully built over thousands of years -- I wish I could say tonight that a lasting peace is in sight.

 

Happily, I can say that war has been avoided. Steady progress toward our ultimate goal has been made. But, so much remains to be done. As a private citizen, I shall never cease to do what little I can to help the world advance along that road.

 

VII.

 

So -- in this my last good night to you as your President -- I thank you for the many opportunities you have given me for public service in war and peace. I trust that in that service you find some things worthy; as for the rest of it, I know you will find ways to improve performance in the future.

 

You and I -- my fellow citizens -- need to be strong in our faith that all nations, under God, will reach the goal of peace with justice. May we be ever unswerving in devotion to principle, confident but humble with power, diligent in pursuit of the Nation's great goals.

 

To all the peoples of the world, I once more give expression to America's prayerful and continuing aspiration:

 

We pray that peoples of all faiths, all races, all nations, may have their great human needs satisfied; that those now denied opportunity shall come to enjoy it to the full; that all who yearn for freedom may experience its spiritual blessings; that those who have freedom will understand, also, its heavy responsibilities; that all who are insensitive to the needs of others will learn charity; that the scourges of poverty, disease and ignorance will be made to disappear from the earth, and that, in the goodness of time, all peoples will come to live together in a peace guaranteed by the binding force of mutual respect and love.

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IN CONGRESS, July 4, 1776.

The unanimous Declaration of the thirteen united States of America,

 

When in the Course of human events, it becomes necessary for one people to dissolve the political bands which have connected them with another, and to assume among the powers of the earth, the separate and equal station to which the Laws of Nature and of Nature's God entitle them, a decent respect to the opinions of mankind requires that they should declare the causes which impel them to the separation.

 

We hold these truths to be self-evident, that all men are created equal, that they are endowed by their Creator with certain unalienable Rights, that among these are Life, Liberty and the pursuit of Happiness.--That to secure these rights, Governments are instituted among Men, deriving their just powers from the consent of the governed, --That whenever any Form of Government becomes destructive of these ends, it is the Right of the People to alter or to abolish it, and to institute new Government, laying its foundation on such principles and organizing its powers in such form, as to them shall seem most likely to effect their Safety and Happiness. Prudence, indeed, will dictate that Governments long established should not be changed for light and transient causes; and accordingly all experience hath shewn, that mankind are more disposed to suffer, while evils are sufferable, than to right themselves by abolishing the forms to which they are accustomed. But when a long train of abuses and usurpations, pursuing invariably the same Object evinces a design to reduce them under absolute Despotism, it is their right, it is their duty, to throw off such Government, and to provide new Guards for their future security.--Such has been the patient sufferance of these Colonies; and such is now the necessity which constrains them to alter their former Systems of Government. The history of the present King of Great Britain is a history of repeated injuries and usurpations, all having in direct object the establishment of an absolute Tyranny over these States. To prove this, let Facts be submitted to a candid world.

 

He has refused his Assent to Laws, the most wholesome and necessary for the public good. He has forbidden his Governors to pass Laws of immediate and pressing importance, unless suspended in their operation till his Assent should be obtained; and when so suspended, he has utterly neglected to attend to them. He has refused to pass other Laws for the accommodation of large districts of people, unless those people would relinquish the right of Representation in the Legislature, a right inestimable to them and formidable to tyrants only. He has called together legislative bodies at places unusual, uncomfortable, and distant from the depository of their public Records, for the sole purpose of fatiguing them into compliance with his measures. He has dissolved Representative Houses repeatedly, for opposing with manly firmness his invasions on the rights of the people. He has refused for a long time, after such dissolutions, to cause others to be elected; whereby the Legislative powers, incapable of Annihilation, have returned to the People at large for their exercise; the State remaining in the meantime exposed to all the dangers of invasion from without, and convulsions within. He has endeavoured to prevent the population of these States; for that purpose obstructing the Laws for Naturalization of Foreigners; refusing to pass others to encourage their migrations hither, and raising the conditions of new Appropriations of Lands. He has obstructed the Administration of Justice, by refusing his Assent to Laws for establishing Judiciary powers. He has made Judges dependent on his Will alone, for the tenure of their offices, and the amount and payment of their salaries. He has erected a multitude of New Offices, and sent hither swarms of Officers to harass our people, and eat out their substance. He has kept among us, in times of peace, Standing Armies without the Consent of our legislatures. He has affected to render the Military independent of and superior to the Civil power. He has combined with others to subject us to a jurisdiction foreign to our constitution, and unacknowledged by our laws; giving his Assent to their Acts of pretended Legislation: For Quartering large bodies of armed troops among us: For protecting them, by a mock Trial, from punishment for any Murders which they should commit on the Inhabitants of these States: For cutting off our Trade with all parts of the world: For imposing Taxes on us without our Consent: For depriving us in many cases, of the benefits of Trial by Jury: For transporting us beyond Seas to be tried for pretended offences For abolishing the free System of English Laws in a neighbouring Province, establishing therein an Arbitrary government, and enlarging its Boundaries so as to render it at once an example and fit instrument for introducing the same absolute rule into these Colonies: For taking away our Charters, abolishing our most valuable Laws, and altering fundamentally the Forms of our Governments: For suspending our own Legislatures, and declaring themselves invested with power to legislate for us in all cases whatsoever. He has abdicated Government here, by declaring us out of his Protection and waging War against us. He has plundered our seas, ravaged our Coasts, burnt our towns, and destroyed the lives of our people. He is at this time transporting large Armies of foreign Mercenaries to complete the works of death, desolation and tyranny, already begun with circumstances of Cruelty & perfidy scarcely paralleled in the most barbarous ages, and totally unworthy the Head of a civilized nation. He has constrained our fellow Citizens taken Captive on the high Seas to bear Arms against their Country, to become the executioners of their friends and Brethren, or to fall themselves by their Hands. He has excited domestic insurrections amongst us, and has endeavoured to bring on the inhabitants of our frontiers, the merciless Indian Savages, whose known rule of warfare, is an undistinguished destruction of all ages, sexes and conditions.

 

In every stage of these Oppressions We have Petitioned for Redress in the most humble terms: Our repeated Petitions have been answered only by repeated injury. A Prince whose character is thus marked by every act which may define a Tyrant, is unfit to be the ruler of a free people.

 

Nor have We been wanting in attentions to our British brethren. We have warned them from time to time of attempts by their legislature to extend an unwarrantable jurisdiction over us. We have reminded them of the circumstances of our emigration and settlement here. We have appealed to their native justice and magnanimity, and we have conjured them by the ties of our common kindred to disavow these usurpations, which, would inevitably interrupt our connections and correspondence. They too have been deaf to the voice of justice and of consanguinity. We must, therefore, acquiesce in the necessity, which denounces our Separation, and hold them, as we hold the rest of mankind, Enemies in War, in Peace Friends.

 

We, therefore, the Representatives of the united States of America, in General Congress, Assembled, appealing to the Supreme Judge of the world for the rectitude of our intentions, do, in the Name, and by Authority of the good People of these Colonies, solemnly publish and declare, That these United Colonies are, and of Right ought to be Free and Independent States; that they are Absolved from all Allegiance to the British Crown, and that all political connection between them and the State of Great Britain, is and ought to be totally dissolved; and that as Free and Independent States, they have full Power to levy War, conclude Peace, contract Alliances, establish Commerce, and to do all other Acts and Things which Independent States may of right do. And for the support of this Declaration, with a firm reliance on the protection of divine Providence, we mutually pledge to each other our Lives, our Fortunes and our sacred Honor.

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“We must hang together, gentlemen...else, we shall most assuredly hang separately.” -- Benjamin Franklin

 

"The tree of liberty must be refreshed from time to time with the blood of patriots and tyrants." -- Thomas Jefferson

 

“When the people fear the government, that's tyranny; when the government fears the people, that's freedom.” -- Thomas Jefferson

  

This image is excerpted from a U.S. GAO report:

www.gao.gov/products/GAO-16-623

 

INFORMATION TECHNOLOGY: Better Management of Interdependencies between Programs Supporting 2020 Census Is Needed

 

Note: The Bureau continues to refine its assumptions as it conducts further research and testing leading up to the 2020 Census.

 

a) The 30 area census offices for identifying addresses where people could live are included in the 250 area census offices planned for the 2020 Census.

 

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

 

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Outlining a Theory of General Creativity . .

. . on a 'Pataphysical projectory

 

Entropy ≥ Memory ● Creativity ²

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published in YSE#22

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La mecanosfera extrae y actualiza configuraciones que existen entre una infinidad de ellas en los campos de la virtualidad. Las máquinas existenciales están al mismo nivel que el ser en su multiplicidad intrínseca. (...) Son para sí mismas su propio material de expresión semiótica. La existencia, como proceso de desterritorialización, es una operación inter-maquínica específica que se superpone a la promoción de intensidades existenciales singularizadas. (...) La existencia no es dialéctica, no es representable. ¡Apenas es vivible!

 

(...) Las máquinas del deseo, las máquinas de la creación estética, en tanto máquinas científicas modifican constantemente nuestras fronteras cósmicas. Como tales, ocupan un lugar eminente dentro de los ensamblajes de subjetivación, ellas mismas llamadas a relevar a nuestras antiguas máquinas sociales que son incapaces de seguir la floración de revoluciones maquinarias que sacuden nuestra época desde todas partes. Más que adoptar una actitud reticente con respecto a la inmensa revolución maquinaria que arrasa el planeta (a riesgo de destruirlo) o agarrarse a sistemas de valores tradicionales, fingiendo reestablecer la trascendencia, el movimiento del progreso, o si se prefiere, el movimiento del proceso, se esforzará por reconciliar los valores con las máquinas.

 

(...) Los sistemas maquínicos se posicionan a sí mismos en un rizoma de interdependencia, situando cada estasis maquínica real en la conjunción de una filiación pasada y un Phylum de futuras mutaciones. Todos los sistemas de valores - religiosos, estéticos, científicos, ecosóficos...- se instalan en esta interfaz maquínica entre lo actual necesario y lo virtual posibilista. (...) De ahí un doble enunciado: finito, territorializado e incorpóreo, infinito.

 

(...) A la estéril oposición entre valor de uso y valor de intercambio es preciso oponer una complexión axiológica que incluya todas las modalidades maquínicas de valoración: los valores del deseo, los valores estéticos, los valores ecológicos, los valores económicos, etc. (...) La ley económica, como la ley jurídica, debe ser deducida del conjunto de Universos del valor, por cuyo colapso lucha constantemente. Su reconstrucción, sobre los escombros desperdigados de las economías planificadas y el neoliberalismo y según nuevas finalidades etico-políticas (ecosofía), requiere, en contraste, una inagotable renovación de la consistencia de los ensamblajes maquínicos de valoración.

 

( Félix Guattari - Caosmosis - 1992 )

 

( traducción por Alicia Pallas )

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rectO-persO | E ≥ m.C² | co~errAnce | TiLt

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

"When we try to pick out anything by itself we find that it is bound fast by a thousand invisible cords that cannot be broken, to everything in the universe. "

—John Muir

 

“A person experiences life as something separated from the rest -a kind of optical delusion of consciousness. Our task must be to free ourselves from this self-imposed prison,and through compassion, to find the reality of Oneness.”

- Albert Einstein

 

“The heart sends blood to every cell of the body, and in this way the cells are nourished. The same blood then flows back to the heart. If the flow is obstructed, the person will die. We need to learn this process of give and take from the heart. For the benefit of others, and also for ourselves, we should have the attitude of caring and sharing. We are all links in the chain of life. If one link is weakened, it will affect the strength of the whole chain.”

—Amma

 

“I think we have to wake up to the fact that everything is connected to everything else. Safety, well-being cannot be individual matters anymore. If others are not safe there is no way that we can be safe. Taking care of others’ safety is at the same time taking care of our own safety. Taking care of others’ well-being is to take care of our own well-being. It is the mind of discrimination and separation that is at the foundation of all violence and hate.”

—Thich Nhat Hanh

 

“An umbilical cord links you to the sun. Another umbilical cord links you to the clouds in the sky. If the clouds were not there, there would be no rain and no water to drink. Without clouds, there is no milk, no tea, no coffee, no ice cream, nothing. There is an umbilical cord linking you to the river; there is one linking you to the forest. If you continue meditating like this, you can see that you are linked to everything and everyone in the cosmos. Your life depends on everything else that exists—on other living beings, but also on plants, minerals, air, water, and earth.”

—Thich Nhat Hanh

  

See in Large...

The artist/event can’t exist without public. They are interdependent. This photo is part of a series about the counterpart of the artist/events: “Spectators’

Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.

 

The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.

 

The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.

 

Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.

 

There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.

 

Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.

 

Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.

 

Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.

 

Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.

 

Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.

 

All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.

 

Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.

 

After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.

 

Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.

 

Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).

 

Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.

 

Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.

 

Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.

 

Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).

 

Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.

 

So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).

 

Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.

 

The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.

 

Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.

 

In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.

 

Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.

 

Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.

 

Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.

 

The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.

 

The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.

 

The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.

 

The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.

 

The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.

 

Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.

 

Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.

 

Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.

 

The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.

 

The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.

 

Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.

 

Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.

 

Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.

 

The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.

 

Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.

 

Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.

 

Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.

 

The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.

 

The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.

 

The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.

 

The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).

 

The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.

 

Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.

 

There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.

 

Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.

 

Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.

 

As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.

 

The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).

 

The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.

 

Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.

 

Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.

 

Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.

 

Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.

 

A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.

 

An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.

  

Yin Yang is a Chinese symbol illustrating "how polar or seemingly contrary forces are interconnected and interdependent in the natural world, and how they give rise to each other in turn." This simple shape of mirrored inverted images is represents many concepts and a variety of religious, and cultural practices including Taoism, the I Ching, Tai Chi and more. Continue reading on Wikipedia.

 

Yin, Yang, slow, fast, soft, hard, yielding, solid, diffuse, focused, cold, hot, wet, dry, passive, aggressive, water, fire, earth, sky, moon, sun, femininity, masculinity, nighttime, daytime.

In the Valley of Parvati....

 

In Hindu mythology, Parvati is Shakti, the wife of Shiva and the gentle aspect of Mahadevi, the Great Goddess.

Shakti from Sanskrit 'shak' - "to be able," meaning sacred force or empowerment, is the primordial cosmic energy and represents the dynamic forces that are thought to move through the entire universe in Hinduism. Shakti is the concept, or personification, of divine feminine creative power, sometimes referred to as 'The Great Divine Mother' in Hinduism. On the earthly plane, Shakti most actively manifests through female embodiment and creativity/fertility, though it is also present in males in its potential, unmanifest form.

 

Not only is the Shakti responsible for creation, it is also the agent of all change. Shakti is cosmic existence as well as liberation, its most significant form being the Kundalini Shakti, a mysterious psychospiritual force. Shakti exists in a state of 'svātantrya', dependence on no-one, being interdependent with the entire universe.

 

Parvati is considered as complete incarnation of 'Adi Parashakti', with all other goddesses being her incarnations or manifestations.

 

"Parvata" is one of the Sanskrit words for "mountain"; "Parvati" translates to "She of the mountains" and refers to Parvati being born the daughter of Himavat, lord of the mountains and the personification of the Himalayas.

 

...and all this divinity you feel when you enter that valley that is named after her.