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Fins are distinctive anatomical features composed of bony
spine Spine or spinal may refer to: Science Biology * Vertebral column, also known as the backbone * Dendritic spine, a small membranous protrusion from a neuron's dendrite * Thorns, spines, and prickles, needle-like structures in plants * Spine (zoolo ...
s or
ray Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gr ...
s protruding from the body of a
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
. They are covered with
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
and joined together either in a webbed fashion, as seen in most
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
, or similar to a flipper, as seen in
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s. Apart from the tail or
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
, fish fins have no direct connection with the
spine Spine or spinal may refer to: Science Biology * Vertebral column, also known as the backbone * Dendritic spine, a small membranous protrusion from a neuron's dendrite * Thorns, spines, and prickles, needle-like structures in plants * Spine (zoolo ...
and are supported only by
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
s. Their principal function is to help the fish swim. Fins located in different places on the fish serve different purposes such as moving forward, turning, keeping an upright position or stopping. Most fish use fins when swimming, flying fish use pectoral fins for gliding, and frogfish use them for crawling. Fins can also be used for other purposes; male
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s and
mosquitofish The western Mosquitofish (''Gambusia affinis'') is a North American freshwater fish, also known commonly, if ambiguously, as simply Mosquitofish or by its generic name, ''Gambusia'', or by the common name gambezi. Its sister species, the easte ...
use a modified fin to deliver sperm,
thresher shark Thresher sharks are large lamniform sharks of the family Alopiidae found in all temperate and tropical oceans of the world; the family contains three extant species, all within the genus ''Alopias''. All three thresher shark species have bee ...
s use their caudal fin to stun prey, reef stonefish have spines in their dorsal fins that inject venom, anglerfish use the first spine of their dorsal fin like a fishing rod to lure prey, and
triggerfish Triggerfish are about 40 species of often brightly colored fish of the family Balistidae. Often marked by lines and spots, they inhabit tropical and subtropical oceans throughout the world, with the greatest species richness in the Indo-Pacif ...
avoid
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s by squeezing into coral crevices and using spines in their fins to lock themselves in place. __TOC__


Types of fins

Fins can either be paired or unpaired. The pectoral and pelvic fins are paired, whereas the dorsal, anal and caudal fins are unpaired and situated along the midline of the body. For every type of fin, there are a number of fish
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
in which this particular fin has been lost during evolution (e.g. pelvic fins in ''
Bobasatrania ''Bobasatrania'' is an extinct genus of prehistoric bony fish that survived the Permian-Triassic extinction event. Fossils of ''Bobasatrania'' were found in beds of Changhsingian (late Permian) to Ladinian (Middle Triassic) age. It was most sp ...
'', caudal fin in
ocean sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, '' Mola alexandrini''. Adults typically weigh between . The sp ...
). In some
clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English te ...
, additional unpaired fins were acquired during evolution (e.g. additional dorsal fins, adipose fin). In some
Acanthodii Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, ...
("spiny sharks"), one or more pairs of "intermediate" or "prepelvic" spines are present between the pectoral and pelvic fins, but these are not associated with fins.


Bony fishes

Bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
es form a taxonomic group called Osteichthyes (or Euteleostomi, which includes also land vertebrates). They have skeletons made of bone mostly, and can be contrasted with
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
es (see below), which have skeletons made mainly of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck ...
(except for their
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
, fin spines, and denticles). Bony fishes are divided into
ray-finned Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or ho ...
and lobe-finned fish. Most living fish are ray-finned, an extremely diverse and abundant group consisting of over 30,000
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. It is the largest class of vertebrates in existence today, making up more than 50% of species. In the distant past, lobe-finned fish were abundant. Nowadays they are show depauperate diversity, with only eight living species. Bony fish have fin spines and rays called lepidotrichia. They typically have
swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their current water depth wit ...
s, which allows the fish to create a neutral balance between sinking and floating without having to use its fins. However, swim bladders are absent in many fish, most notably in Lungfishes, which are the only fish to have retained the primitive lung present in the common ancestor of bony fish from which swim bladders evolved. Bony fishes also have an operculum, which helps them breathe without having to use fins to swim.


Lobe-fins

Lobe-finned fishes form a class of
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
es called Sarcopterygii. They have fleshy,
lobed The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
, paired fins, which are joined to the body by a series of bones. The fins of lobe-finned fish differ from those of all other fish in that each is borne on a fleshy, lobe-like, scaly stalk extending from the body. Pectoral and pelvic fins have articulations resembling those of tetrapod limbs. These fins evolved into legs of the first
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
land vertebrates (
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s) in the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
Period. Sarcopterygians also possess two dorsal fins with separate bases, as opposed to the single dorsal fin of most
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
(except some teleosts). The
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
is either heterocercal (only fossil taxa) or diphycercal. '' Latimeria'' is a lobe-finned fish which is still extant. There are two species, the West Indian Ocean coelacanth (''Latimeria chalumnae'') and the Indonesian coelacanth (''Latimeria menadoensis''). Coelacanths are thought to have evolved into roughly their current form about 408 million years ago, during the early Devonian. Locomotion of the coelacanths is unique to their kind. To move around, coelacanths most commonly take advantage of up or downwellings of the current and drift. They use their paired fins to stabilize their movement through the water. While on the ocean floor their paired fins are not used for any kind of movement. Coelacanths can create thrust for quick starts by using their caudal fins. Due to the high number of fins they possess, coelacanths have high maneuverability and can orient their bodies in almost any direction in the water. They have been seen doing headstands and swimming belly up. It is thought that their rostral organ helps give the coelacanth electroperception, which aids in their movement around obstacles.
Lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
are also living lobe-finned fish. They occur in Africa ('' Protopterus''), Australia ('' Neoceratodus''), and South America (''
Lepidosiren The South American lungfish (''Lepidosiren paradoxa'') is the single species of lungfish found in swamps and slow-moving waters of the Amazon, Paraguay, and lower Paraná River basins in South America. Notable as an obligate air-breather, it is t ...
''). Lungfish evolved during the Devonian Period. Genetic studies and paleontological data confirm that lungfish are the closest living relatives of land vertebrates. Fin arrangement and body shape is relatively conservative in lobe-finned fishes. However, there are a few examples from the fossil record that show aberrant morphologies, such as ''
Allenypterus ''Allenypterus'' is a genus of a prehistoric lobe-finned fish which lived during the Bashkirian stage of the Late Carboniferous period, 318 million years ago). Fossils have been discovered in Bear Gulch Limestone The Bear Gulch Limestone is ...
'', '' Rebellatrix'', ''
Foreyia ''Foreyia'' is an extinct genus of coelacanth lobe-finned fish which lived during the Middle Triassic period in what is now Canton of Graubünden, Switzerland. It contains a single species ''F. maxkuhni''. Description and classification ''F. ma ...
'' or the tetrapodomorphs.


Diversity of fins in lobe-finned fishes


Ray-fins

Ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
es form a class of
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
es called Actinopterygii. Their fins contain spines or rays. A fin may contain only spiny rays, only soft rays, or a combination of both. If both are present, the spiny rays are always anterior. Spines are generally stiff and sharp. Rays are generally soft, flexible, segmented, and may be branched. This segmentation of rays is the main difference that separates them from spines; spines may be flexible in certain species, but they will never be segmented. Spines have a variety of uses. In
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
, they are used as a form of defense; many catfish have the ability to lock their spines outwards.
Triggerfish Triggerfish are about 40 species of often brightly colored fish of the family Balistidae. Often marked by lines and spots, they inhabit tropical and subtropical oceans throughout the world, with the greatest species richness in the Indo-Pacif ...
also use spines to lock themselves in crevices to prevent them being pulled out. Lepidotrichia are usually composed of
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
, but in early osteichthyans such as '' Cheirolepis'', there was also dentine and enamel. They are segmented and appear as a series of disks stacked one on top of another. They may have been derived from dermal scales. The genetic basis for the formation of the fin rays is thought to be genes coded for the production of certain proteins. It has been suggested that the evolution of the tetrapod limb from lobe-finned fishes is related to the loss of these proteins.


Diversity of fins in ray-finned fishes


Cartilaginous fishes

Cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
es form a class of fishes called Chondrichthyes. They have skeletons made of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck ...
rather than
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
. The class includes
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s,
ray Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gr ...
s and
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
s. Shark fin skeletons are elongated and supported with soft and unsegmented rays named ceratotrichia, filaments of elastic protein resembling the horny
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up Scale (anatomy), scales, hair, Nail ...
in hair and feathers. Originally the pectoral and pelvic girdles, which do not contain any dermal elements, did not connect. In later forms, each pair of fins became ventrally connected in the middle when scapulocoracoid and puboischiadic bars evolved. In rays, the pectoral fins have connected to the head and are very flexible. One of the primary characteristics present in most sharks is the heterocercal tail, which aids in locomotion. Most sharks have eight fins. Sharks can only drift away from objects directly in front of them because their fins do not allow them to move in the tail-first direction. Unlike modern cartilaginous fish, members of
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
chondrichthyan lineages (e.g. the climatiids and the diplacanthids) possessed pectoral dermal plates as well as dermal spines associated with the paired fins. The oldest species demonstrating these features is the acanthodian '' Fanjingshania renovata'' from the lower
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
( Aeronian) of China. Fanjingshania possess compound pectoral plates composed of dermal scales fused to a bony plate and fin spines formed entirely of bone. Fin spines associated with the dorsal fins are rare among extant cartilaginous fishes, but are present, for instance, in '' Heterodontus'' or ''
Squalus ''Squalus'' is a genus of dogfish sharks in the family Squalidae. Commonly known as spurdogs, these sharks are characterized by smooth dorsal fin spines, teeth in upper and lower jaws similar in size, caudal peduncle with lateral keels; upper ...
''. Dorsal fin spines are typically developed in many fossil groups, such as in Hybodontiformes,
Ctenacanthiformes Ctenacanthiformes is an extinct order of chondrichthyan fish. They possessed ornamented fin spines and cladodont dentition. Members of the family Ctenacanthidae may have survived into the Cretaceous based on teeth found in deep water deposits ...
or Xenacanthida. In '' Stethacanthus'', the first dorsal fin spine was modified, forming a spine-brush complex. As with most fish, the tails of sharks provide thrust, making speed and acceleration dependent on tail shape.
Caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
shapes vary considerably between shark species, due to their evolution in separate environments. Sharks possess a
heterocercal Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
caudal fin in which the dorsal portion is usually noticeably larger than the ventral portion. This is because the shark's
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
extends into that dorsal portion, providing a greater surface area for
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
attachment. This allows more efficient locomotion among these negatively buoyant cartilaginous fish. By contrast, most bony fish possess a homocercal caudal fin. Tiger sharks have a large upper lobe, which allows for slow cruising and sudden bursts of speed. The tiger shark must be able to twist and turn in the water easily when hunting to support its varied diet, whereas the porbeagle shark, which hunts schooling fish such as mackerel and herring, has a large lower lobe to help it keep pace with its fast-swimming prey. Other tail adaptations help sharks catch prey more directly, such as the
thresher shark Thresher sharks are large lamniform sharks of the family Alopiidae found in all temperate and tropical oceans of the world; the family contains three extant species, all within the genus ''Alopias''. All three thresher shark species have bee ...
's usage of its powerful, elongated upper lobe to stun fish and squid. On the other hand, rays rely on their enlarged pectoral fins for propulsion. Similarly enlarged pectoral fins can be found in the
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
Petalodontiformes (e.g. ''
Belantsea ''Belantsea'' (named after a legendary ancestor of the Crow Nation) is a genus of extinct petalodontid cartilaginous fish that lived during the Lower Carboniferous, about 350 million years ago. Its fossils are found in the Bear Gulch Limestone ...
'', ''
Janassa ''Janassa'' is an extinct genus of petalodont cartilaginous fish that lived in marine environments in what is now central United States of America and Europe during the Carboniferous and upper Permian. It is known from teeth and a few poorl ...
'', '' Menaspis''), which belong to Holocephali (ratfish and their fossil relatives), or in ''
Aquilolamna ''Aquilolamna'' is an extinct genus of shark-like elasmobranch from the Late Cretaceous (Turonian)-aged Agua Nueva Formation of Mexico. It is currently known to contain only one species, ''A. milarcae'', also known as the eagle shark, and it is c ...
'' ( Selachimorpha) and ''
Squatinactis ''Squatinactis'' is a genus of extinct elasmobranch Chondrichthyes known from the Carboniferous aged Bear gulch limestone in Montana. This fish was discovered in 1974 by Richard Lund. The type specimen, named CMNH 46133, consists of a brain case ...
'' (Squatinactiformes). Some cartilaginous fishes have an eel-like locomotion (e.g. ''
Chlamydoselachus ''Chlamydoselachus'' is a genus of sharks and the sole extant member of the family Chlamydoselachidae, in the order Hexanchiformes. It contains two extant and several extinct species. The most widely known species still surviving is the frilled s ...
'', ''
Thrinacoselache ''Thrinacodus'' is an extinct genus of basal elasmobranch, found worldwide from the Late Devonian-Lower Carboniferous. Most species are only known from their tricuspid teeth. ''T. gracia'', originally placed in the separate genus ''Thrinacoselach ...
'', '' Phoebodus'')


Diversity of fins in cartilaginous fishes


Shark finning

According to the Humane Society International, approximately 100 million sharks are killed each year for their fins, in an act known as shark finning. After the fins are cut off, the mutilated sharks are thrown back in the water and left to die. In some countries of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, shark fins are a culinary delicacy, such as
shark fin soup Shark fin soup is a traditional soup or stewed dish served in parts of China, Taiwan, and Southeast Asia. The shark fins provide texture, while the taste comes from the other soup ingredients. It is commonly served at special occasions such as ...
. Currently, international concerns over the
sustainability Specific definitions of sustainability are difficult to agree on and have varied in the literature and over time. The concept of sustainability can be used to guide decisions at the global, national, and individual levels (e.g. sustainable livi ...
and
welfare Welfare, or commonly social welfare, is a type of government support intended to ensure that members of a society can meet basic human needs such as food and shelter. Social security may either be synonymous with welfare, or refer specifical ...
of sharks have impacted consumption and availability of shark fin soup worldwide. Shark finning is prohibited in many countries.


Fin functions


Generating thrust

Foil shaped fins generate
thrust Thrust is a reaction force described quantitatively by Newton's third law. When a system expels or accelerates mass in one direction, the accelerated mass will cause a force of equal magnitude but opposite direction to be applied to that ...
when moved, the lift of the fin sets water or air in motion and pushes the fin in the opposite direction. Aquatic animals get significant
thrust Thrust is a reaction force described quantitatively by Newton's third law. When a system expels or accelerates mass in one direction, the accelerated mass will cause a force of equal magnitude but opposite direction to be applied to that ...
by moving fins back and forth in water. Often the tail fin is used, but some aquatic animals generate thrust from pectoral fins.
Cavitation Cavitation is a phenomenon in which the static pressure of a liquid reduces to below the liquid's vapour pressure, leading to the formation of small vapor-filled cavities in the liquid. When subjected to higher pressure, these cavities, ca ...
occurs when negative pressure causes bubbles (cavities) to form in a liquid, which then promptly and violently collapse. It can cause significant damage and wear.Franc, Jean-Pierre and Michel, Jean-Marie (2004
''Fundamentals of Cavitation''
Springer. .
Cavitation damage can occur to the tail fins of powerful swimming marine animals, such as dolphins and tuna. Cavitation is more likely to occur near the surface of the ocean, where the ambient water pressure is relatively low. Even if they have the power to swim faster, dolphins may have to restrict their speed because collapsing cavitation bubbles on their tail are too painful. Cavitation also slows tuna, but for a different reason. Unlike dolphins, these fish do not feel the bubbles, because they have bony fins without nerve endings. Nevertheless, they cannot swim faster because the cavitation bubbles create a vapor film around their fins that limits their speed. Lesions have been found on tuna that are consistent with cavitation damage. Scombrid fishes (tuna, mackerel and bonito) are particularly high-performance swimmers. Along the margin at the rear of their bodies is a line of small rayless, non-retractable fins, known as
finlet Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
s. There has been much speculation about the function of these finlets. Research done in 2000 and 2001 by Nauen and Lauder indicated that "the finlets have a hydrodynamic effect on local flow during steady swimming" and that "the most posterior finlet is oriented to redirect flow into the developing tail vortex, which may increase thrust produced by the tail of swimming mackerel". Fish use multiple fins, so it is possible that a given fin can have a hydrodynamic interaction with another fin. In particular, the fins immediately upstream of the caudal (tail) fin may be proximate fins that can directly affect the flow dynamics at the caudal fin. In 2011, researchers using volumetric imaging techniques were able to generate "the first instantaneous three-dimensional views of wake structures as they are produced by freely swimming fishes". They found that "continuous tail beats resulted in the formation of a linked chain of vortex rings" and that "the dorsal and anal fin wakes are rapidly entrained by the caudal fin wake, approximately within the timeframe of a subsequent tail beat".


Controlling motion

Once motion has been established, the motion itself can be controlled with the use of other fins. The bodies of reef fishes are often shaped differently from open water fishes. Open water fishes are usually built for speed, streamlined like torpedoes to minimise friction as they move through the water. Reef fish operate in the relatively confined spaces and complex underwater landscapes of
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
s. For this manoeuvrability is more important than straight line speed, so coral reef fish have developed bodies which optimize their ability to dart and change direction. They outwit predators by dodging into fissures in the reef or playing hide and seek around coral heads.Alevizon WS (1994
"Pisces Guide to Caribbean Reef Ecology"
Gulf Publishing Company
The pectoral and pelvic fins of many reef fish, such as
butterflyfish The butterflyfish are a group of conspicuous tropical marine fish of the family Chaetodontidae; the bannerfish and coralfish are also included in this group. The approximately 129 species in 12 genera are found mostly on the reefs of the Atlan ...
, damselfish and
angelfish Angelfish may refer to: *Several groups of fish: **Freshwater angelfish, tropical cichlids of the genus ''Pterophyllum'' **Marine angelfish of the family Pomacanthidae **Atlantic pomfret (''Brama brama''), sold by fishmongers as "angelfish" in Sou ...
, have evolved so they can act as brakes and allow complex manoeuvres.Ichthyology
''Florida Museum of Natural History''. Retrieved 22 November 2012.
Many reef fish, such as
butterflyfish The butterflyfish are a group of conspicuous tropical marine fish of the family Chaetodontidae; the bannerfish and coralfish are also included in this group. The approximately 129 species in 12 genera are found mostly on the reefs of the Atlan ...
, damselfish and
angelfish Angelfish may refer to: *Several groups of fish: **Freshwater angelfish, tropical cichlids of the genus ''Pterophyllum'' **Marine angelfish of the family Pomacanthidae **Atlantic pomfret (''Brama brama''), sold by fishmongers as "angelfish" in Sou ...
, have evolved bodies which are deep and laterally compressed like a pancake, and will fit into fissures in rocks. Their pelvic and pectoral fins have evolved differently, so they act together with the flattened body to optimise manoeuvrability. Some fishes, such as puffer fish,
filefish The filefish (Monacanthidae) are a diverse family of tropical to subtropical tetraodontiform marine fish, which are also known as foolfish, leatherjackets or shingles. They live in the Atlantic, Pacific and Indian Oceans. Filefish are closely r ...
and trunkfish, rely on pectoral fins for swimming and hardly use tail fins at all.


Reproduction

Male
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
es (sharks and rays), as well as the males of some live-bearing ray finned fishes, have fins that have been modified to function as
intromittent organ An intromittent organ is any external organ of a male organism that is specialized to deliver sperm during copulation. Intromittent organs are found most often in terrestrial species, as most non-mammalian aquatic species fertilize their eggs e ...
s, reproductive appendages which allow internal fertilization. In ray finned fish they are called ''gonopodia'' or ''andropodia'', and in cartilaginous fish they are called ''claspers''. ''Gonopodia'' are found on the males of some species in the Anablepidae and
Poeciliidae The Poeciliidae are a family of freshwater fishes of the order Cyprinodontiformes, the tooth-carps, and include well-known live-bearing aquarium fish, such as the guppy, molly, platy, and swordtail. The original distribution of the family was ...
families. They are anal fins that have been modified to function as movable intromittent organs and are used to impregnate females with milt during mating. The third, fourth and fifth rays of the male's anal fin are formed into a tube-like structure in which the sperm of the fish is ejected. When ready for mating, the gonopodium becomes erect and points forward towards the female. The male shortly inserts the organ into the sex opening of the female, with hook-like adaptations that allow the fish to grip onto the female to ensure impregnation. If a female remains stationary and her partner contacts her vent with his gonopodium, she is fertilized. The sperm is preserved in the female's oviduct. This allows females to fertilize themselves at any time without further assistance from males. In some species, the gonopodium may be half the total body length. Occasionally the fin is too long to be used, as in the "lyretail" breeds of '' Xiphophorus helleri''. Hormone treated females may develop gonopodia. These are useless for breeding. Similar organs with similar characteristics are found in other fishes, for example the ''andropodium'' in the '' Hemirhamphodon'' or in the Goodeidae or the ''gonopodium'' in the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma an ...
''
Saurichthys ''Saurichthys'' (from el, σαῦρος , 'lizard' and el, ἰχθῦς 'fish') is an extinct genus of predatory ray-finned fish from the Triassic period. It type genus family Saurichthyidae (Changhsingian-Middle Jurassic), and the largest ...
'', the oldest known example of
viviparity Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the ...
in a ray-finned fish. '' Claspers'' are found on the males of
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
es. They are the posterior part of the pelvic fins that have also been modified to function as intromittent organs, and are used to channel semen into the female's cloaca during copulation. The act of mating in sharks usually includes raising one of the claspers to allow water into a siphon through a specific orifice. The clasper is then inserted into the cloaca, where it opens like an umbrella to anchor its position. The siphon then begins to contract expelling water and sperm.


Other functions

Other uses of fins include walking and perching on the sea floor, gliding over water, cooling of body temperature, stunning of prey, display (scaring of predators, courtship), defence (venomous fin spines, locking between corals), luring of prey, and attachment structures. The
Indo-Pacific sailfish The Indo-Pacific sailfish (''Istiophorus platypterus'') is a sailfish native to the Indian and Pacific Oceans and is naturalized in the Atlantic where it has entered the Mediterranean Sea via the Suez Canal as a Lessepsian migrant. It is dark b ...
has a prominent dorsal fin. Like scombroids and other billfish, they streamline themselves by retracting their dorsal fins into a groove in their body when they swim. The huge dorsal fin, or sail, of the sailfish is kept retracted most of the time. Sailfish raise them if they want to herd a school of small fish, and also after periods of high activity, presumably to cool down.''Aquatic Life of the World''
pp. 332–333, Marshall Cavendish Corporation, 2000. .
The oriental flying gurnard has large pectoral fins which it normally holds against its body, and expands when threatened to scare predators. Despite its name, it is a
demersal fish Demersal fish, also known as groundfish, live and feed on or near the bottom of seas or lakes (the demersal zone).Walrond Carl . "Coastal fish - Fish of the open sea floor"Te Ara - the Encyclopedia of New Zealand. Updated 2 March 2009 They oc ...
, not a flying fish, and uses its pelvic fins to walk along the bottom of the ocean. Fins can have an adaptive significance as sexual ornaments. During courtship, the female
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted th ...
, '' Pelvicachromis taeniatus'', displays a large and visually arresting purple pelvic fin. "The researchers found that males clearly preferred females with a larger pelvic fin and that pelvic fins grew in a more disproportionate way than other fins on female fish."


Evolution


Evolution of paired fins

There are two prevailing hypotheses that have been historically debated as models for the evolution of paired fins in fish: the gill arch theory and the lateral fin-fold theory. The former, commonly referred to as the “ Gegenbaur hypothesis,” was posited in 1870 and proposes that the “paired fins are derived from gill structures”. This fell out of popularity in favor of the lateral fin-fold theory, first suggested in 1877, which proposes that paired fins budded from longitudinal, lateral folds along the epidermis just behind the gills. There is weak support for both hypotheses in the fossil record and in embryology. However, recent insights from developmental patterning have prompted reconsideration of both theories in order to better elucidate the origins of paired fins.


Classical theories

Karl Gegenbaur's concept of the “Archipterygium” was introduced in 1876. It was described as a gill ray, or “joined cartilaginous stem,” that extended from the gill arch. Additional rays arose from along the arch and from the central gill ray. Gegenbaur suggested a model of transformative homology – that all vertebrate paired fins and limbs were transformations of the Archipterygium. Based on this theory, paired appendages such as pectoral and pelvic fins would have differentiated from the branchial arches and migrated posteriorly. However, there has been limited support for this hypothesis in the fossil record both morphologically and phylogenically. In addition, there was little to no evidence of an anterior-posterior migration of pelvic fins. Such shortcomings of the gill-arch theory led to its early demise in favor of the lateral fin-fold theory proposed by St. George Jackson Mivart, Francis Balfour, and James Kingsley Thacher. The lateral fin-fold theory hypothesized that paired fins developed from lateral folds along the body wall of the fish. Just as segmentation and budding of the median fin fold gave rise to the median fins, a similar mechanism of fin bud segmentation and elongation from a lateral fin fold was proposed to have given rise to the paired pectoral and pelvic fins. However, there was little evidence of a lateral fold-to-fin transition in the fossil record. In addition, it was later demonstrated phylogenically that pectoral and pelvic fins arise from distinct evolutionary and mechanistic origins.


Evolutionary developmental biology

Recent studies in the ontogeny and evolution of paired appendages have compared finless vertebrates – such as lampreys – with
chondricthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
, the most basal living vertebrate with paired fins. In 2006, researchers found that the same genetic programming involved in the segmentation and development of median fins was found in the development of paired appendages in catsharks. Although these findings do not directly support the lateral fin-fold hypothesis, the original concept of a shared median-paired fin evolutionary developmental mechanism remains relevant. A similar renovation of an old theory may be found in the developmental programming of chondricthyan gill arches and paired appendages. In 2009, researchers at the
University of Chicago The University of Chicago (UChicago, Chicago, U of C, or UChi) is a private research university in Chicago, Illinois. Its main campus is located in Chicago's Hyde Park neighborhood. The University of Chicago is consistently ranked among the b ...
demonstrated that there are shared molecular patterning mechanisms in the early development of the chondricthyan gill arch and paired fins. Findings such as these have prompted reconsideration of the once-debunked gill-arch theory.


From fins to limbs

Fish are the ancestors of all mammals, reptiles, birds and amphibians. In particular, terrestrial
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s (four-legged animals) evolved from fish and made their first forays onto land 400 million years ago. They used paired pectoral and pelvic fins for locomotion. The pectoral fins developed into forelegs (arms in the case of humans) and the pelvic fins developed into hind legs. Much of the genetic machinery that builds a walking limb in a tetrapod is already present in the swimming fin of a fish. In 2011, researchers at
Monash University Monash University () is a public research university based in Melbourne, Victoria, Australia. Named for prominent World War I general Sir John Monash, it was founded in 1958 and is the second oldest university in the state. The university has ...
in Australia used primitive but still living
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
"to trace the evolution of pelvic fin muscles to find out how the load-bearing hind limbs of the tetrapods evolved." Further research at the University of Chicago found bottom-walking lungfishes had already evolved characteristics of the walking gaits of terrestrial tetrapods. In a classic example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
, the pectoral limbs of
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s,
birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
and
bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bir ...
further evolved along independent paths into flying wings. Even with flying wings there are many similarities with walking legs, and core aspects of the genetic blueprint of the pectoral fin have been retained. The first mammals appeared during the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
period (between 298.9 and 252.17 million years ago). Several groups of these mammals started returning to the sea, including the
cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
s (whales, dolphins and porpoises). Recent DNA analysis suggests that cetaceans evolved from within the even-toed ungulates, and that they share a common ancestor with the
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two exta ...
. About 23 million years ago another group of bearlike land mammals started returning to the sea. These were the
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
. What had become walking limbs in cetaceans and seals evolved independently into new forms of swimming fins. The forelimbs became
flippers Flipper may refer to: Common meanings *Flipper (anatomy), a forelimb of an aquatic animal, useful for steering and/or propulsion in water *Alternate name for a swimfin, footwear that boosts human swimming efficiency * Flipper (cricket), a type of ...
, while the hindlimbs were either lost (cetaceans) or also modified into flipper (pinnipeds). In cetaceans, the tail gained two fins at the end, called a
fluke Fluke may refer to: Biology * Fluke (fish), a species of marine flatfish * Fluke (tail), the lobes of the tail of a cetacean, such as dolphins or whales, ichthyosaurs, mosasaurs, plesiosaurs, and metriorhynchids. * Fluke (flatworm), parasiti ...
. Fish tails are usually vertical and move from side to side. Cetacean flukes are horizontal and move up and down, because cetacean spines bend the same way as in other mammals.
Ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...
s are ancient reptiles that resembled dolphins. They first appeared about 245 million years ago and disappeared about 90 million years ago.
"This sea-going reptile with terrestrial ancestors converged so strongly on fishes that it actually evolved a
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through c ...
and tail fin for improved aquatic locomotion. These structures are all the more remarkable because they evolved from nothing — the ancestral terrestrial reptile had no hump on its back or blade on its tail to serve as a precursor."Martill D.M. (1993). "Soupy Substrates: A Medium for the Exceptional Preservation of Ichthyosaurs of the Posidonia Shale (Lower Jurassic) of Germany". ''Kaupia - Darmstädter Beiträge zur Naturgeschichte'', 2 : 77-97.
The biologist
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Goul ...
said the ichthyosaur was his favorite example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Fins or flippers of varying forms and at varying locations (limbs, body, tail) have also evolved in a number of other tetrapod groups, including diving birds such as penguins (modified from wings), sea turtles (forelimbs modified into flippers), mosasaurs (limbs modified into flippers), and sea snakes (vertically expanded, flattened tail fin).


Robotic fins

The use of fins for the propulsion of aquatic animals can be remarkably effective. It has been calculated that some fish can achieve a
propulsive A prokinetic agent (also gastroprokinetic agent, gastrokinetic agent or propulsive) is a type of drug which enhances gastrointestinal motility by increasing the frequency or strength of contractions, but without disrupting their rhythm. They are ...
efficiency greater than 90%. Fish can accelerate and maneuver much more effectively than
boat A boat is a watercraft of a large range of types and sizes, but generally smaller than a ship, which is distinguished by its larger size, shape, cargo or passenger capacity, or its ability to carry boats. Small boats are typically found on i ...
s or
submarine A submarine (or sub) is a watercraft capable of independent operation underwater. It differs from a submersible, which has more limited underwater capability. The term is also sometimes used historically or colloquially to refer to remotely op ...
, and produce less water disturbance and noise. This has led to biomimetic studies of underwater robots which attempt to emulate the locomotion of aquatic animals. An example is the Robot Tuna built by th
Institute of Field Robotics
to analyze and mathematically model thunniform motion. In 2005, the Sea Life London Aquarium displayed three robotic fish created by the computer science department at the University of Essex. The fish were designed to be autonomous, swimming around and avoiding obstacles like real fish. Their creator claimed that he was trying to combine "the speed of tuna, acceleration of a pike, and the navigating skills of an eel." The ''AquaPenguin'', developed by Festo of Germany, copies the streamlined shape and propulsion by front flippers of
penguin Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adap ...
s. Festo also developed ''AquaRay'', ''AquaJelly'' and ''AiraCuda'', respectively emulating the locomotion of manta rays, jellyfish and barracuda. In 2004, Hugh Herr at MIT prototyped a
biomechatronic Biomechatronics is an applied interdisciplinary science that aims to integrate biology and mechatronics (electrical, electronics, and mechanical engineering). It also encompasses the fields of robotics and neuroscience. Biomechatronic devices encom ...
robotic fish with a living
actuator An actuator is a component of a machine that is responsible for moving and controlling a mechanism or system, for example by opening a valve. In simple terms, it is a "mover". An actuator requires a control device (controlled by control signal) a ...
by surgically transplanting muscles from frog legs to the robot and then making the robot swim by pulsing the muscle fibers with electricity. Robotic fish offer some research advantages, such as the ability to examine an individual part of a fish design in isolation from the rest of the fish. However, this risks oversimplifying the biology so key aspects of the animal design are overlooked. Robotic fish also allow researchers to vary a single parameter, such as flexibility or a specific motion control. Researchers can directly measure forces, which is not easy to do in live fish. "Robotic devices also facilitate three-dimensional kinematic studies and correlated hydrodynamic analyses, as the location of the locomotor surface can be known accurately. And, individual components of a natural motion (such as outstroke vs. instroke of a flapping appendage) can be programmed separately, which is certainly difficult to achieve when working with a live animal."


See also

* Cephalopod fin * Fin and flipper locomotion * Fish locomotion * Polydactyly in early tetrapods * RoboTuna *
Shark fin soup Shark fin soup is a traditional soup or stewed dish served in parts of China, Taiwan, and Southeast Asia. The shark fins provide texture, while the taste comes from the other soup ingredients. It is commonly served at special occasions such as ...
*
Tradeoffs for locomotion in air and water Certain species of fish and birds are able to locomote in both air and water, two fluid media with very different properties. A fluid is a particular phase of matter that deforms under shear stresses and includes any type of liquid or gas. Becau ...
* Undulatory locomotion


References


Citations


Bibliography

*


Further reading

* Hall, Brian K (2007
''Fins into Limbs: Evolution, Development, and Transformation''
University of Chicago Press. . * Helfman G, Collette BB, Facey DE and Bowen BW (2009
"Functional morphology of locomotion and feeding"
Chapter 8, pp. 101–116. In:''The Diversity of Fishes: Biology'', John Wiley & Sons. . * *


External links


Homology of fin lepidotrichia in osteichthyan fishes


Earthlife Web

''HowStuffWorks''. Accessed 30 January 2012. {{Authority control Fish anatomy