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A fish scale is a small rigid plate that grows out of the
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 cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
of a fish. The skin of most
jawed fish Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...
es is covered with these protective scales, which can also provide effective
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
through the use of
reflection Reflection or reflexion may refer to: Science and technology * Reflection (physics), a common wave phenomenon ** Specular reflection, reflection from a smooth surface *** Mirror image, a reflection in a mirror or in water ** Signal reflection, in ...
and colouration, as well as possible hydrodynamic advantages. The term ''scale'' derives from the
Old French Old French (, , ; Modern French: ) was the language spoken in most of the northern half of France from approximately the 8th to the 14th centuries. Rather than a unified language, Old French was a linkage of Romance dialects, mutually intelligib ...
, meaning a shell pod or husk. Scales vary enormously in size, shape, structure, and extent, ranging from strong and rigid armour plates in fishes such as
shrimpfish Shrimpfish, also called razorfish, are five small species of marine fishes in the subfamily Centriscinae of the family Centriscidae. The species in the genera ''Aeoliscus'' and ''Centriscus'' are found in relatively shallow tropical parts of the ...
es and
boxfish Ostraciidae is a family of squared, bony fish belonging to the order Tetraodontiformes, closely related to the pufferfishes and filefishes. Fish in the family are known variously as boxfishes, cofferfishes, cowfishes and trunkfishes. It contains ...
es, to microscopic or absent in fishes such as
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s and
anglerfish The anglerfish are fish of the teleost order Lophiiformes (). They are bony fish named for their characteristic mode of predation, in which a modified luminescent fin ray (the esca or illicium) acts as a lure for other fish. The luminescence ...
es. The
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
of a scale can be used to identify the species of fish it came from. Scales originated within the jawless
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and th ...
s, ancestors to all jawed fishes today. Most bony fishes are covered with the cycloid scales of
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
and
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
, or the ctenoid scales of
perch Perch is a common name for fish of the genus ''Perca'', freshwater gamefish belonging to the family Percidae. The perch, of which three species occur in different geographical areas, lend their name to a large order of vertebrates: the Percif ...
, or the ganoid scales of
sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...
s and
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s.
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 (
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s and
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
) are covered with placoid scales. Some species are covered instead by
scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s, and others have no outer covering on part or all of the skin. Fish scales are part of the fish's
integumentary system The integumentary system is the set of organs forming the outermost layer of an animal's body. It comprises the skin and its appendages, which act as a physical barrier between the external environment and the internal environment that it serves ...
, and are produced from the
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...
layer of the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
, which distinguishes them from
reptile scale Reptile skin is covered with scutes or scales which, along with many other characteristics, distinguish reptiles from animals of other classes. They are made of alpha and beta-keratin and are formed from the epidermis (contrary to fish, in which t ...
s. The same
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s involved in tooth and hair development in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s are also involved in scale development. The placoid scales of cartilaginous fishes are also called dermal denticles and are structurally homologous with vertebrate teeth. Most fish are also covered in a layer of
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is ...
or slime which can protect against pathogens such as bacteria, fungi, and viruses, and reduce surface resistance when the fish swims.


Thelodont scales

The bony scales of
thelodont Thelodonti (from Greek: "feeble teeth")Maisey, John G., Craig Chesek, and David Miller. Discovering fossil fishes. New York: Holt, 1996. is a class (biology), class of extinct jawless fishes with distinctive Fish scale, scales instead of large pl ...
s, the most abundant form of
fossil fish The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish ...
, are well understood. The scales were formed and shed throughout the organisms' lifetimes, and quickly separated after their death. Bone, a tissue that is both resistant to mechanical damage and relatively prone to fossilization, often preserves internal detail, which allows the
histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
and growth of the scales to be studied in detail. The scales comprise a non-growing "crown" composed of
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
, with a sometimes-ornamented
enameloid Enameloid, also known as durodentine or vitrodentine, is an Tooth enamel, enamel-like Tissue (biology), tissue found in many fish. It is the primary outer component of shark Odontode, odontodes (teeth and dermal denticles). Although the origin of e ...
upper surface and an aspidine base. Its growing base is made of cell-free bone, which sometimes developed anchorage structures to fix it in the side of the fish. Beyond that, there appear to be five types of bone-growth, which may represent five natural groupings within the thelodonts—or a spectrum ranging between the end members meta- (or ortho-) dentine and mesodentine tissues. Each of the five scale morphs appears to resemble the scales of more derived groupings of fish, suggesting that thelodont groups may have been stem groups to succeeding clades of fish. However, using scale morphology alone to distinguish species has some pitfalls. Within each organism, scale shape varies hugely according to body area, with intermediate forms appearing between different areas—and to make matters worse, scale morphology may not even be constant within one area. To confuse things further, scale morphologies are not unique to taxa, and may be indistinguishable on the same area of two different species. The morphology and histology of thelodonts provides the main tool for quantifying their diversity and distinguishing between species, although ultimately using such convergent traits is prone to errors. Nonetheless, a framework comprising three groups has been proposed based upon scale morphology and histology. Comparisons to modern shark species have shown that thelodont scales were functionally similar to those of modern cartilaginous fish, and likewise has allowed an extensive comparison between ecological niches.


Cosmoid scales

Cosmoid scales are found only on ancient
lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
es, including some of the earliest
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
es (subclass
Dipnoi 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 ...
), and in Crossopterygii, including the living
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
in a modified form (see elasmoid scales, below). They were probably derived from a fusion of placoid-ganoid scales. The inner part of the scales is made of dense
lamellar A ''lamella'' (plural ''lamellae'') is a small plate or flake, from the Latin, and may also be used to refer to collections of fine sheets of material held adjacent to one another, in a gill-shaped structure, often with fluid in between though s ...
bone called isopedine. On top of this lies a layer of spongy or
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
bone supplied with blood vessels, followed by a complex
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
-like layer called
cosmine Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales. Description As traditionally des ...
with a superficial outer coating of
vitrodentine Enameloid, also known as durodentine or vitrodentine, is an enamel-like tissue found in many fish. It is the primary outer component of shark odontodes (teeth and dermal denticles). Although the origin of enameloid is debated, it is probably ho ...
. The upper surface is
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 scales, hair, nails, feathers, ho ...
. Cosmoid scales increase in size through the growth of the lamellar bone layer.


Elasmoid scales

Elasmoid scales are thin, imbricated scales composed of a layer of dense, lamellar collagen bone called isopedine, above which is a layer of tubercles usually composed of bone, as in ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
''. The layer of dentine that was present in the first lobe-finned fish is usually reduced, as in the extant
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
, or entirely absent, as in extant
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 ...
and in the Devonian ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
''. Elasmoid scales have appeared several times over the course of fish evolution. They are present in some
lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
es, such as all extant and some extinct
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
es, as well as the
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
s which have modified cosmoid scales that lack cosmine and are thinner than true cosmoid scales. They are also present in some tetrapodomorphs like ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
'', amiids, and teleosts, whose cycloid and ctenoid scales represent the least mineralized elasmoid scales. The
zebrafish The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family ( Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ...
elasmoid scales are used in the lab to study bone mineralization process, and can be cultured (kept) outside of the organism.


Ganoid scales

Ganoid scales are found in the
sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...
s,
paddlefish Paddlefish (family Polyodontidae) are a family of ray-finned fish belonging to order Acipenseriformes, and one of two living groups of the order alongside sturgeons (Acipenseridae). They are distinguished from other fish by their titular elongla ...
es,
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s,
bowfin The bowfin (''Amia calva'') is a bony fish, native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being the sole surviving species of the Halecomorphi ...
, and
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
s. They are derived from cosmoid scales and often have serrated edges. They are covered with a layer of hard enamel-like
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
in the place of
cosmine Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales. Description As traditionally des ...
, and a layer of inorganic bone salt called ''
ganoine Ganoine or ''ganoin'' is a glassy, often multi-layered mineralized tissue that covers the scales, cranial bones and fin rays in some non-teleost ray-finned fishes, such as gars, bichirs, and coelacanths. It is composed of rod-like, pseudoprism ...
'' in place of
vitrodentine Enameloid, also known as durodentine or vitrodentine, is an enamel-like tissue found in many fish. It is the primary outer component of shark odontodes (teeth and dermal denticles). Although the origin of enameloid is debated, it is probably ho ...
. Ganoine is a characteristic component of ganoid scales. It is a glassy, often multi-layered mineralized tissue that covers the scales, as well as the
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
bones and fin rays in some non-teleost
ray-finned fishes 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 hor ...
, such as
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s,
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
s, and
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
s. It is composed of rod-like
apatite Apatite is a group of phosphate minerals, usually hydroxyapatite, fluorapatite and chlorapatite, with high concentrations of OH−, F− and Cl− ions, respectively, in the crystal. The formula of the admixture of the three most common e ...
crystallites. Ganoine is an ancient feature of ray-finned fishes, being found for example on the scales of
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
actinopteryigian ''
Cheirolepis ''Cheirolepis'' (from el, χείρ , 'hand' and el, λεπίς 'scale') is an extinct genus of ray-finned fish that lived in the Devonian period of Europe and North America. It is the only genus yet known within the family Cheirolepidae and th ...
''. While often considered a synapomorphic character of ray-finned fishes, ganoine or ganoine-like tissues are also found on the extinct
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, which ...
. It has been suggested ganoine is homologous to
tooth enamel Tooth enamel is one of the four major Tissue (biology), tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the Crown (tooth), crown. The ...
in vertebrates or even considered a type of enamel. Most ganoid scales are
rhomboidal Traditionally, in two-dimensional geometry, a rhomboid is a parallelogram in which adjacent sides are of unequal lengths and angles are non-right angled. A parallelogram with sides of equal length (equilateral) is a rhombus but not a rhomboid. ...
(diamond-shaped) and connected by peg-and-socket joints. They are usually thick and fit together more like a jigsaw rather than overlapping like other scales. In this way, ganoid scales are nearly impenetrable and are excellent protection against predation. File:Alligator gar fish.jpg, The
alligator gar The alligator gar (''Atractosteus spatula'') is a ray-finned euryhaline fish related to the bowfin in the infraclass Holostei . It is the largest species in the gar family, and among the largest freshwater fish in North America. The fossil ...
has a tough armouring of
rhomboidal Traditionally, in two-dimensional geometry, a rhomboid is a parallelogram in which adjacent sides are of unequal lengths and angles are non-right angled. A parallelogram with sides of equal length (equilateral) is a rhombus but not a rhomboid. ...
-shaped ganoid scales. File:Acipenser oxyrhynchus (recropped).png, The
sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...
has rows of ganoid scales enlarged into scute-like armour plates. File:Bowfin fish image.jpg, The ganoid scales on a
bowfin The bowfin (''Amia calva'') is a bony fish, native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being the sole surviving species of the Halecomorphi ...
are reduced in size and resemble cycloid scales.
In sturgeons, the scales are greatly enlarged into armour plates along the sides and back, while in the bowfin the scales are greatly reduced in thickness to resemble cycloid scales. File:Earrings from the ganoid scales of alligator gar.jpg, Earrings made from the ganoid scales of an alligator gar File:Schmelzschuppenfischkonkretion.jpg, Fossil of a primitive rayfin with ganoid scales File:Fish Fossil (FindID 64765).jpg, Ganoid scales on a fossilised ''
Lepidotes ''Lepidotes'' (from el, λεπιδωτός , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has been considered a wastebasket taxon, characterised by "general features, such as thi ...
'', circa. 130 mya
Native Americans and people of the Caribbean used the tough ganoid scales of the
alligator gar The alligator gar (''Atractosteus spatula'') is a ray-finned euryhaline fish related to the bowfin in the infraclass Holostei . It is the largest species in the gar family, and among the largest freshwater fish in North America. The fossil ...
for arrow heads, breastplates, and as shielding to cover plows. In current times jewellery is made from these scales.


Leptoid scales

Leptoid (bony-ridge) scales are found on higher-order bony fish, the
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s (the more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
clade 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 term, ...
of ray-finned fishes). The outer part of these scales fan out with bony ridges while the inner part is criss-crossed with fibrous connective tissue. Leptoid scales are thinner and more translucent than other types of scales, and lack the hardened enamel-like or dentine layers. Unlike ganoid scales, further scales are added in concentric layers as the fish grows. Leptoid scales overlap in a head-to-tail configuration, like roof tiles, making them more flexible than cosmoid and ganoid scales. This arrangement allows a smoother flow of water over the body, and reduces drag. The scales of some species exhibit bands of uneven seasonal growth called ''annuli'' (singular ''annulus''). These bands can be used to age the fish. Leptoid scales come in two forms: cycloid (smooth) and ctenoid (comb-like).


Cycloid scales

Cycloid (circular) scales have a smooth texture and are uniform, with a smooth outer edge or margin. They are most common on fish with soft fin rays, such as
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
and
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
.


Ctenoid scales

Ctenoid (toothed) scales are like cycloid scales, except they have small teeth or
spinule Spinules are small spines or thorns (vertebral columns) that are part of biological and manmade structures. The word originates from the Latin word and is often used in botany and zoology. The presence or absence of spinules, and their shape, can ...
s called ctenii along their outer or posterior edges. Because of these teeth, the scales have a rough texture. They are usually found on fishes with spiny fin rays, such as the perch-like fishes. These scales contain almost no bone, being composed of a surface layer containing
hydroxyapatite Hydroxyapatite, also called hydroxylapatite (HA), is a naturally occurring mineral form of calcium apatite with the formula Ca5(PO4)3(OH), but it is usually written Ca10(PO4)6(OH)2 to denote that the crystal unit cell comprises two entities. ...
and calcium carbonate and a deeper layer composed mostly of
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
. The enamel of the other scale types is reduced to superficial ridges and ctenii. Ctenoid scales, similar to other epidermal structures, originate from placodes and distinctive cellular differentiation makes them exclusive from other structures that arise from the
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
.Kawasaki, Kenta C., "A Genetic Analysis of Cichlid Scale Morphology" (2016). Masters Theses May 2014 - current. 425. http://scholarworks.umass.edu/masters_theses_2/425 Development starts near 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 ...
, along the
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
of the fish. The development process begins with an accumulation of
fibroblast A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s between the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
and
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
.
Collagen fibrils Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
begin to organize themselves in the dermal layer, which leads to the initiation of mineralization. The circumference of the scales grows first, followed by thickness when overlapping layers mineralize together. Ctenoid scales can be further subdivided into three types: * Crenate scales, where the margin of the scale bears indentations and projections. * Spinoid scales, where the scale bears spines that are continuous with the scale itself. * True ctenoid scales, where the spines on the scale are distinct structures. Most ray-finned fishes have ctenoid scales. Some species of
flatfish A flatfish is a member of the Ray-finned fish, ray-finned demersal fish order (biology), order Pleuronectiformes, also called the Heterosomata, sometimes classified as a suborder of Perciformes. In many species, both eyes lie on one side of the ...
es have ctenoid scales on the eyed side and cycloid scales on the blind side, while other species have ctenoid scales in males and cycloid scales in females.


Reflection

Many teleost fish are covered with highly reflective scales which function as small mirrors and give the appearance of silvered glass. Reflection through silvering is widespread or dominant in fish of the open sea, especially those that live in the top 100 metres. A transparency effect can be achieved by silvering to make an animal's body highly reflective. At medium depths at sea, light comes from above, so a mirror oriented vertically makes animals such as fish invisible from the side. The
marine hatchetfish Marine hatchetfishes or deep-sea hatchetfishes are small deep-sea mesopelagic ray-finned fish of the stomiiform subfamily Sternoptychinae. They should not be confused with the freshwater hatchetfishes, which are not particularly closely related T ...
is extremely flattened laterally (side to side), leaving the body just millimetres thick, and the body is so silvery as to resemble
aluminium foil Aluminium foil (or aluminum foil in North American English; often informally called tin foil) is aluminium prepared in thin metal leaves with a thickness less than ; thinner gauges down to are also commonly used. Standard household foil is typ ...
. The mirrors consist of microscopic structures similar to those used to provide
structural coloration Structural coloration in animals, and a few plants, is the production of colour by microscopically structured surfaces fine enough to interfere with visible light instead of pigments, although some structural coloration occurs in combination wit ...
: stacks of between 5 and 10 crystals of
guanine Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine (uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is called ...
spaced about ¼ of a wavelength apart to interfere constructively and achieve nearly 100 per cent reflection. In the deep waters that the hatchetfish lives in, only blue light with a wavelength of 500 nanometres percolates down and needs to be reflected, so mirrors 125 nanometres apart provide good camouflage. Most fish in the upper ocean are camouflaged by silvering. In fish such as the
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Oceans, i ...
, which lives in shallower water, the mirrors must reflect a mixture of wavelengths, and the fish accordingly has crystal stacks with a range of different spacings. A further complication for fish with bodies that are rounded in cross-section is that the mirrors would be ineffective if laid flat on the skin, as they would fail to reflect horizontally. The overall mirror effect is achieved with many small reflectors, all oriented vertically. Fish scales with these properties are used in some cosmetics, since they can give a shimmering effect to makeup and lipstick.


Placoid scales

Placoid (pointed, tooth-shaped) scales are found in the
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:
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s,
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
. They are also called dermal denticles. Placoid scales are structurally homologous with
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
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, tear ...
("denticle" translates to "small tooth"), having a central pulp cavity supplied with
blood vessel The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away ...
s, surrounded by a conical layer of
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
, all of which sits on top of a rectangular basal plate that rests on the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
. The outermost layer is composed of
vitrodentine Enameloid, also known as durodentine or vitrodentine, is an enamel-like tissue found in many fish. It is the primary outer component of shark odontodes (teeth and dermal denticles). Although the origin of enameloid is debated, it is probably ho ...
, a largely inorganic enamel-like substance. Placoid scales cannot grow in size, but rather more scales are added as the fish increases in size. Similar scales can also be found under the head of the denticle herring. The amount of scale coverage is much less in rays. Rhomboidal scales with the properties of both placoid and ganoid scales are suspected to exist in modern jawed fish ancestors: jawless
ostracoderms Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and ...
and then jawed
placoderms Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally 'plate-skinned') is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were ...
.


Shark skin

Shark skin is almost entirely covered by small placoid scales. The scales are supported by spines, which feel rough when stroked in a backward direction, but when flattened by the forward movement of water, create tiny
vortices In fluid dynamics, a vortex ( : vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in th ...
that reduce
hydrodynamic In physics and engineering, fluid dynamics is a subdiscipline of fluid mechanics that describes the flow of fluids—liquids and gases. It has several subdisciplines, including ''aerodynamics'' (the study of air and other gases in motion) and ...
drag and reduce
turbulence In fluid dynamics, turbulence or turbulent flow is fluid motion characterized by chaotic changes in pressure and flow velocity. It is in contrast to a laminar flow, which occurs when a fluid flows in parallel layers, with no disruption between ...
, making swimming both more efficient and quieter compared to that of bony fishes. It also serves a role in anti-fouling by exhibiting the
lotus effect The lotus effect refers to self-cleaning properties that are a result of ultrahydrophobicity as exhibited by the leaves of ''Nelumbo'', the lotus flower. Dirt particles are picked up by water droplets due to the micro- and nanoscopic architect ...
. All denticles are composed of an interior pulp cavity with a nervous and arterial supply rooted in the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
to supply the denticle with mucus. Denticles contain riblet structures that protrude from the surface of the scale; under a microscope this riblet can look like a hook or ridges coming out of the scale. The overall shape of the protrusion from the denticle is dependent on the type of shark and can be generally described with two appearances. The first is a scale in which ridges are placed laterally down the shark and parallel with the flow of the water. The second form is a smooth scale with what looks like a hooked riblet curling out of the surface aiming towards the posterior side of the shark. Both riblet shapes assist in creating a turbulent
boundary layer In physics and fluid mechanics, a boundary layer is the thin layer of fluid in the immediate vicinity of a bounding surface formed by the fluid flowing along the surface. The fluid's interaction with the wall induces a no-slip boundary condi ...
forcing the
laminar flow In fluid dynamics, laminar flow is characterized by fluid particles following smooth paths in layers, with each layer moving smoothly past the adjacent layers with little or no mixing. At low velocities, the fluid tends to flow without lateral mi ...
farther away from the sharks skin. Unlike bony fish, sharks have a complicated dermal corset made of flexible
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
ous
fiber Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...
s arranged as a
helical Helical may refer to: * Helix, the mathematical concept for the shape * Helical engine, a proposed spacecraft propulsion drive * Helical spring, a coilspring * Helical plc, a British property company, once a maker of steel bar stock * Helicoil A t ...
network surrounding their body. The corset works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy. Depending on the position of these placoid scales on the body, they can be flexible and can be passively erected, allowing them to change their angle of attack. These scales also have riblets which are aligned in the direction of flow, these riblets reduce the drag force acting on the shark skin by pushing the vortex further away from the skin surface, inhibiting any high-velocity cross-stream flow.


Scale morphology

The general anatomy of the scales varies, but all of them can be divided into three parts: the crown, the neck and the base. The scale pliability is related to the size of the base of the scale. The scales with higher flexibility have a smaller base, and thus are less rigidly attached to the ''stratum laxum.'' On the crown of the fast-swimming sharks there are a series of parallel riblets or ridges which run from an anterior to posterior direction. Analyzing the three components of the scale it can be concluded that the base of the denticle does not come into contact with any portion of the fluid flow. The crown and the neck of the denticles however play a key role and are responsible for creating the turbulent vortices and
eddies In fluid dynamics, an eddy is the swirling of a fluid and the reverse current created when the fluid is in a turbulent flow regime. The moving fluid creates a space devoid of downstream-flowing fluid on the downstream side of the object. Fluid beh ...
found near the skin's surface. Because denticles come in so many different shapes and sizes, it can be expected that not all shapes will produce the same type of
turbulent flow In fluid dynamics, turbulence or turbulent flow is fluid motion characterized by chaotic changes in pressure and flow velocity. It is in contrast to a laminar flow, which occurs when a fluid flows in parallel layers, with no disruption between ...
. During a recent research experiment
biomimetic Biomimetics or biomimicry is the emulation of the models, systems, and elements of nature for the purpose of solving complex human problems. The terms "biomimetics" and "biomimicry" are derived from grc, βίος (''bios''), life, and μίμησ ...
samples of shark denticles with a crescent like microstructure and tested in a water tank using a traction table as a slide. The experiment showed that the surface with denticles experienced a 10% drag reduction overall versus the smooth sample. The reason for this drag reduction was that the turbulent vortices became trapped between the denticles, creating a ‘cushion like’ barrier against the laminar flow. This same type of experiment was performed by another research group which implemented more variation in their biomimetic sample. The second group arrived at the same conclusion as the first, however, because their experiment contained more variation within the samples they were able to achieve a high degree of experimental accuracy. In conclusion they stated that more practical shapes were more durable than ones with intricate ridge-lines. The practical shapes were low profile and contained trapezoidal or semi-circular trough-like cross sections, and were less effective but nonetheless reduced drag by 6 or 7%.


Drag reduction

Sharks decrease drag and overall
cost of transport The energy cost of transport quantifies the Energy efficiency in transportation, energy efficiency of transporting an animal or vehicle from one place to another. As a dimensionless quantity, it allows for the comparison of dissimilar animals or mod ...
(COT) through multiple different avenues.
Pressure drag Parasitic drag, also known as profile drag, is a type of aerodynamic drag that acts on any object when the object is moving through a fluid. Parasitic drag is a combination of form drag and skin friction drag. It affects all objects regardless of ...
is created from the pressure difference between the anterior and posterior side of the shark due to the amount of volume that is pushed past the shark to propel itself forward. This type of drag is also directly proportional to the
laminar flow In fluid dynamics, laminar flow is characterized by fluid particles following smooth paths in layers, with each layer moving smoothly past the adjacent layers with little or no mixing. At low velocities, the fluid tends to flow without lateral mi ...
. When the laminar flow increases around the fish the pressure drag does as well. Frictional drag is a result of the interaction between the fluid against the shark's skin and can vary depending on how the boundary layer changes against the surface of the fish. The riblets impede the cross-stream translation of the streamwise vortices in the viscous sublayer. The mechanism is complex and not yet understood fully. Basically, the riblets inhibit the vortex formation near the surface because the vortex cannot fit in the valleys formed by the riblets. This pushes the vortex further up from the surface, interacting only with the riblet tips, not causing any high-velocity flow in the valleys. Since this high velocity flow now only interacts with the riblet-tip, which is a very small surface area, the momentum transfer which causes drag is now much lower than before, thereby effectively reducing drag. Also, this reduces the cross-stream velocity fluctuations, which aids in momentum transfer too. Recent research has shown that there is a pre and post breakdown regime in the near-wall boundary layer where the sublayer thickens at a declining rate and then abruptly undergoes a breakdown into turbulent vortices before finally collapsing. This system is completely self regulating and mediates the growth and decay cycle; the vortices accumulate during the growth period and abruptly liquidated in into Strouhal arrays of hairpin vortices lifting off the wall. Lifting vortices are what push the boundary layer out and away from the surface of the shark which results in reducing the overall drag experienced by the fish. The rough,
sandpaper upright=1.35, Sheets of sandpaper with different grit sizes (40 (coarse), 80, 150, 240, 600 (fine)). Sandpaper and glasspaper are names used for a type of coated abrasive that consists of sheets of paper or cloth with abrasive material glued to ...
-like texture of shark and ray skin, coupled with its toughness, has led it to be valued as a source of rawhide
leather Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and hogs, ...
, called
shagreen Shagreen is a type of rawhide consisting of rough untanned skin, historically from a horse's or onager's back, or from shark or ray. Etymology The word derives from the French ''chagrin'' and is related to Italian ''zigrino'' and Venetian '' ...
. One of the many historical applications of shark shagreen was in making hand-grips for
sword A sword is an edged, bladed weapon intended for manual cutting or thrusting. Its blade, longer than a knife or dagger, is attached to a hilt and can be straight or curved. A thrusting sword tends to have a straighter blade with a pointed ti ...
s. The rough texture of the skin is also used in
Japanese cuisine Japanese cuisine encompasses the regional and traditional foods of Japan, which have developed through centuries of political, economic, and social changes. The traditional cuisine of Japan (Japanese: ) is based on rice with miso soup and other ...
to make
grater A grater, also known as a shredder, is a kitchen utensil used to grate foods into fine pieces. Uses Food preparation Several types of graters feature different sizes of grating slots, and can therefore aid in the preparation of a variety of ...
s called '' oroshiki'', by attaching pieces of shark skin to wooden boards. The small size of the scales grates the food very finely.


Technical application

In the marine industry, there is an extremely large market and need for anti-fouling surfaces. In laymen's terms, fouling is known as the process by which something becomes encrusted with material from the surrounding environment such as
barnacle A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in eros ...
s,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, and green sludge. Dermal denticles are an extremely promising area of research for this type of application due to the fact that sharks are among the only fish without build up or growth on their scales. Studies by the
U.S. Navy The United States Navy (USN) is the maritime service branch of the United States Armed Forces and one of the eight uniformed services of the United States. It is the largest and most powerful navy in the world, with the estimated tonnage o ...
have shown that if a biomimetic material can be engineered, it could potentially lead to fuel cost savings for military vessels of up to 45%. There are many examples of
biomimetic material Biomimetic materials are materials developed using inspiration from nature. This may be useful in the design of composite materials. Natural structures have inspired and innovated human creations. Notable examples of these natural structures includ ...
s and surfaces based on the structure of aquatic organisms, including sharks. Such applications intend to enable more efficient movement through fluid mediums such as air, water, and oil. Surfaces that mimic the skin of sharks have also been used in order to keep microorganisms and
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
from coating the hulls of submarines and ships. One variety is traded as " sharklet". A lot of the new methods for replicating shark skin involve the use of
polydimethylsiloxane Polydimethylsiloxane (PDMS), also known as dimethylpolysiloxane or dimethicone, belongs to a group of polymeric organosilicon compounds that are commonly referred to as silicones. PDMS is the most widely used silicon-based organic polymer, as its ...
(PDMS) for creating a mold. Usually the process involves taking a flat piece of shark skin, covering it with the PDMS to form a mold and pouring PDMS into that mold again to get a shark skin replica. This method has been used to create a biomimetic surface which has
superhydrophobic Ultrahydrophobic (or superhydrophobic) surfaces are highly hydrophobic, i.e., extremely difficult to wet. The contact angles of a water droplet on an ultrahydrophobic material exceed 150°. This is also referred to as the lotus effect, after the ...
properties, exhibiting the
lotus effect The lotus effect refers to self-cleaning properties that are a result of ultrahydrophobicity as exhibited by the leaves of ''Nelumbo'', the lotus flower. Dirt particles are picked up by water droplets due to the micro- and nanoscopic architect ...
. One study found that these biomimetic surfaces reduced drag by up to 9%, while with flapping motion drag reduction reached 12.3%. Along with marine applications, the aerospace industry can also benefit from these biomimetic designs.
Parametric model In statistics, a parametric model or parametric family or finite-dimensional model is a particular class of statistical models. Specifically, a parametric model is a family of probability distributions that has a finite number of parameters. Def ...
ing has been done on shark denticles with a wide range of design variations such as low and high-profile vortex generators. These biomimetic models were designed and analyzed to see the effects of applying the denticle-like structures to the wings of various airplanes. During the simulation, it was noted that the sample altered how the low and high
angles of attack In fluid dynamics, angle of attack (AOA, α, or \alpha) is the angle between a reference line on a body (often the chord line of an airfoil) and the vector representing the relative motion between the body and the fluid through which it is m ...
reacted. Out of both the low and high-profile samples tested, the low profile vortex generators outperformed the current smooth wing structures by 323%. This increase in performance is due to a separation bubble in the denticles wake and stream-wise vortices that replenish momentum lost in the boundary layer due to skin friction.


Scutes

Scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s are similar to scales and serve the same function. Unlike the scales of fish, which are formed from the epidermis, scutes are formed in the lower vascular layer of the skin and the epidermal element is only the top surface. Forming in the living dermis, the scutes produce a horny outer layer, that is superficially similar to that of scales. Scute comes from Latin for ''shield'', and can take the form of: * an external shield-like bony plate, or * a modified, thickened scale that often is keeled or spiny, or * a projecting, modified (rough and strongly ridged) scale, usually associated with the lateral line, or on the caudal peduncle forming caudal keels, or along the ventral profile. Some fish, such as
pineconefish Pinecone fishes are small and unusual marine fish of the family ''Monocentridae''. The family contains just four species in two genera, one of which is monotypic. Their distribution is limited to tropical and subtropical waters of the Indo-Pac ...
, are completely or partially covered in scutes. River herrings and
threadfin Threadfins are silvery grey perciform fish of the family Polynemidae. Found in tropical to subtropical waters throughout the world, the threadfin family contains eight genera and about 40 species. An unrelated species sometimes known by the name ...
s have an abdominal row of scutes, which are scales with raised, sharp points that are used for protection. Some jacks have a row of scutes following the
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
on either side.


Scale development

Scales typically appear late in the development of fish. In the case of
zebrafish The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family ( Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ...
, it takes 30 days after fertilization before the different layers needed to start forming the scales have differentiated and become organized. For this it is necessary that consolidation of the
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...
occurs, then
morphogenesis Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of devel ...
is induced, and finally the process of differentiation or late
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
occurs. * Mesenchyme consolidation: The consolidation or structuring of the mesenchyme originates during the development of the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
. This process depends on whether the fish is cartilaginous or bony. For cartilaginous fish the structuring originates through the formation of two layers. The first is superficial and wide and the second is thin and compact. These two layers are separated by
mesenchymal cell Mesenchymal stem cells (MSCs) also known as mesenchymal stromal cells or medicinal signaling cells are multipotent stromal cells that can differentiate into a variety of cell types, including osteoblasts (bone cells), chondrocytes (cartilage cel ...
s. Bony fish generate an
acellular Non-cellular life, or acellular life is life that exists without a cellular structure for at least part of its life cycle. Historically, most (descriptive) definitions of life postulated that an organism must be composed of one or more cells, ...
substrate organized by perpendicularly by
collagen fiber Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whol ...
s. Subsequently, for both fish the
fibroblast A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s elongate. These penetrate the compact layer of the mesenchyme, which consolidates prior to the formation of the scale, in order to initiate the dermal plate. * Morphogenesis induction: The morphogenesis is due to the formation of the epidermal
papilla Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...
, which is generated by joining the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
and
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
through a process of
invagination Invagination is the process of a surface folding in on itself to form a cavity, pouch or tube. In developmental biology, invagination is a mechanism that takes place during gastrulation. This mechanism or cell movement happens mostly in the vegetal ...
. Morphogenesis begins at the time when fibroblasts are relocated to the upper part of the compact mesenchyme. Throughout this process, the basal cells of the
epithelium Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...
form a delimiting layer, which is located in the upper part of the mesenchyme. Subsequently, these cells will differentiate in the area where the scale
primordium A primordium (; plural: primordia; synonym: anlage) in embryology, is an organ or tissue in its earliest recognizable stage of development. Cells of the primordium are called primordial cells. A primordium is the simplest set of cells capable o ...
will arise. * Differentiation or late metamorphosis: This differentiation is generated by two different forms according to the type of scale being formed. The formation of elasmoid scales (cycloids and ctenoids) occurs through the formation of a space between the matrix of the epidermal papilla. This space contains collagen fibers. Around this space elasmoblasts differentiate and are responsible for generating the necessary material for the formation of the scale. Subsequently, matrix mineralization occurs, allowing the scale to acquire the rigid characteristic that identifies them. Unlike elasmoid scales, ganoid scales are composed of mineralized and non-mineralized collagen in different regions. The formation of these occurs through the entry of the surface cells of the mesenchyme into the matrix, the latter is composed of collagen fibers and is located around the vascular capillaries, thus giving rise to vascular cavities. At this point, elasmoblasts are replaced by
osteoblast Osteoblasts (from the Greek language, Greek combining forms for "bone", ὀστέο-, ''osteo-'' and βλαστάνω, ''blastanō'' "germinate") are cell (biology), cells with a single Cell nucleus, nucleus that synthesize bone. However, in the p ...
s, thus forming bone. The patches of the matrix of the scale that are not
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes ...
are composed of compacted collagen that allow it to maintain the union with the mesenchyme. This are known as
Sharpey fibers Sharpey's fibres (bone fibres, or perforating fibres) are a matrix of connective tissue consisting of bundles of strong predominantly type I collagen fibres connecting periosteum to bone. They are part of the outer fibrous layer of periosteum, ...
. One of the genes that regulate the development of scale formation in fish is the
sonic hedgehog Sonic hedgehog protein (SHH) is encoded for by the ''SHH'' gene. The protein is named after the character ''Sonic the Hedgehog''. This signaling molecule is key in regulating embryonic morphogenesis in all animals. SHH controls organogenesis and ...
(shh) gene, which by means of the (shh) protein, involved in
organogenesis Organogenesis is the phase of embryonic development that starts at the end of gastrulation and continues until birth. During organogenesis, the three germ layers formed from gastrulation (the ectoderm, endoderm, and mesoderm) form the internal orga ...
and in the process of
cellular communication A mobile phone, cellular phone, cell phone, cellphone, handphone, hand phone or pocket phone, sometimes shortened to simply mobile, cell, or just phone, is a portable telephone that can make and receive calls over a radio frequency link whil ...
, enable the formation of the scales. The
apolipoprotein E Apolipoprotein E (APOE) is a protein involved in the metabolism of fats in the body of mammals. A subtype is implicated in Alzheimer's disease and cardiovascular disease. APOE belongs to a family of fat-binding proteins called apolipoproteins. ...
(ApoE), that allows the transport and metabolism of
triglyceride A triglyceride (TG, triacylglycerol, TAG, or triacylglyceride) is an ester derived from glycerol and three fatty acids (from ''tri-'' and ''glyceride''). Triglycerides are the main constituents of body fat in humans and other vertebrates, as w ...
s and
cholesterol Cholesterol is any of a class of certain organic molecules called lipids. It is a sterol (or modified steroid), a type of lipid. Cholesterol is biosynthesized by all animal cells and is an essential structural component of animal cell mem ...
, has an interaction with shh, because ApoE provides cholesterol to the shh signaling pathway. It has been shown that during the process of
cell differentiation Cellular differentiation is the process in which a stem cell alters from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellula ...
and
interaction Interaction is action that occurs between two or more objects, with broad use in philosophy and the sciences. It may refer to: Science * Interaction hypothesis, a theory of second language acquisition * Interaction (statistics) * Interactions o ...
, the level of ApoE transcription is high, which has led to the conclusion that this protein is important for the late development of scales.


Modified scales

Different groups of fish have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
a number of modified scales to serve various functions. * Almost all fishes have a
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
, a system of
mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, ...
s that detect water movements. In bony fishes, the scales along the lateral line have central pores that allow water to contact the sensory cells. * The dorsal fin spines of
dogfish shark Squalidae, more commonly known as dogfish, dog sharks, or spiny dogfish, are one of several families of sharks categorized under Squaliformes, making it the second largest order of sharks, numbering 119 species across 7 families. Having earned t ...
s and chimaeras, the stinging tail spines of
stingrays Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae (sixgill stingray), Plesiobatidae ...
, and the "saw" teeth of
sawfish Sawfish, also known as carpenter sharks, are a family of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined with sharp transverse teeth, arranged in a way that resembles a saw. They are among the largest fish wi ...
es and
sawshark A sawshark or saw shark is a member of a shark order (Pristiophoriformes ) bearing a unique long, saw-like Rostrum (anatomy), rostrum (snout or bill) edged with sharp teeth, which they use to slash and disable their prey. There are eight species ...
s are fused and modified placoid scales. *
Surgeonfish Acanthuridae are the family of surgeonfishes, tangs, and unicornfishes. The family includes about 86 extant species of marine fish living in tropical seas, usually around coral reefs. Many of the species are brightly colored and popular in ...
have a scalpel-like blade, which is a modified scale, on either side of the
caudal peduncle 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 ...
. * Some
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Oceans, i ...
s,
anchovies An anchovy is a small, common forage fish of the family Engraulidae. Most species are found in marine waters, but several will enter brackish water, and some in South America are restricted to fresh water. More than 140 species are placed in 1 ...
, and
halfbeak Hemiramphidae is a family of fishes that are commonly called halfbeaks, spipe fish or spipefish. They are a geographically widespread and numerically abundant family of epipelagic fish inhabiting warm waters around the world. The halfbeaks are ...
s have deciduous scales, which are easily shed and aid in escaping predators. * Male '' Percina'' darters have a row of enlarged caducous scales between the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two en ...
s and the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
. *
Porcupine fish Porcupinefish are fish belonging to the family Diodontidae (order Tetraodontiformes), also commonly called blowfish and, sometimes, balloonfish and globefish. They are sometimes collectively called pufferfish, not to be confused with the morpholo ...
es have scales modified into large external spines. * By contrast, pufferfish have thinner, more hidden spines than porcupine fish, which become visible only when the fish puffs up. Unlike the porcupine fish, these spines are not modified scales, but develop under the control of the same network of genes that produce feathers and hairs in other vertebrates. File:FMIB 45807 Atopomycterus nichthemerus.jpeg,
Porcupine fish Porcupinefish are fish belonging to the family Diodontidae (order Tetraodontiformes), also commonly called blowfish and, sometimes, balloonfish and globefish. They are sometimes collectively called pufferfish, not to be confused with the morpholo ...
have scales modified into spines. File:Tetrodon patoca Achilles 182.jpg, Pufferfish spines are not modified scales but are developed by an independent gene network.


Fish without scales

File:Synchiropus splendidus 2 Luc Viatour cropped.png, Mandarinfish lack scales and protect themselves with a layer of smelly and bitter slime. Fish without scales usually evolve alternatives to the protection scales can provide, such as tough leathery skin or bony plates. *
Jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostomes ...
(
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s and
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
es) have smooth skin without scales and without dermal bone. Lampreys get some protection from a tough leathery skin. Hagfish exude copious quantities slime or
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is ...
if they are threatened. They can tie themselves in an
overhand knot The overhand knot is one of the most fundamental knots, and it forms the basis of many others, including the simple noose, overhand loop, angler's loop, reef knot, fisherman's knot, Half hitch, and water knot. The overhand knot is a stopper, e ...
, scraping off the slime as they go and freeing themselves from a predator. * Most
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s are scaleless, though some species are covered with tiny smooth cycloid scales. * Most
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, ...
lack scales, though several families have body armour in the form of dermal plates or some sort of scute. * Mandarinfish lack scales and have a layer of smelly and bitter slime which blocks out disease and probably discourages predators, implying their bright coloration is
aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
. *
Anglerfish The anglerfish are fish of the teleost order Lophiiformes (). They are bony fish named for their characteristic mode of predation, in which a modified luminescent fin ray (the esca or illicium) acts as a lure for other fish. The luminescence ...
have loose, thin skin often covered with fine forked dermal prickles or
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s, but they do not have regular scales. They rely on camouflage to avoid the attention of predators, while their loose skin makes it difficult for predators to grab them. Many groups of bony fishes, including
pipefish Pipefishes or pipe-fishes (Syngnathinae) are a subfamily of small fishes, which, together with the seahorses and seadragons (''Phycodurus'' and ''Phyllopteryx''), form the family Syngnathidae. Description Pipefish look like straight-bodied seah ...
,
seahorse A seahorse (also written ''sea-horse'' and ''sea horse'') is any of 46 species of small marine fish in the genus ''Hippocampus''. "Hippocampus" comes from the Ancient Greek (), itself from () meaning "horse" and () meaning "sea monster" or " ...
s,
boxfish Ostraciidae is a family of squared, bony fish belonging to the order Tetraodontiformes, closely related to the pufferfishes and filefishes. Fish in the family are known variously as boxfishes, cofferfishes, cowfishes and trunkfishes. It contains ...
,
poachers Poaching has been defined as the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set a ...
, and several families of
stickleback The sticklebacks are a family of ray-finned fishes, the Gasterosteidae which have a Holarctic distribution in fresh, brackish and marine waters. They were thought to be related to the pipefish and seahorses but are now thought to be more closel ...
s, have developed external bony plates, structurally resembling placoid scales, as protective armour against predators. * Seahorses lack scales but have thin skin stretched over a bony plate armour arranged in rings through the length of their bodies. * In boxfish, the plates fuse together to form a rigid shell or
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
enclosing the entire body. These bony plates are not modified scales but skin that has been ossified. Because of this heavy armour boxfish are limited to slow movements, but few other fish are able to eat the adults. File:Lactoria fornasini1.jpg,
Boxfish Ostraciidae is a family of squared, bony fish belonging to the order Tetraodontiformes, closely related to the pufferfishes and filefishes. Fish in the family are known variously as boxfishes, cofferfishes, cowfishes and trunkfishes. It contains ...
have plates of ossified skin fused together to form a rigid shell. File:Hippocampus guttulatus Achilles 174.jpg,
Seahorse A seahorse (also written ''sea-horse'' and ''sea horse'') is any of 46 species of small marine fish in the genus ''Hippocampus''. "Hippocampus" comes from the Ancient Greek (), itself from () meaning "horse" and () meaning "sea monster" or " ...
s have thin skin stretched over bony plates arranged in rings.
Some fish, such as hoki and swordfish, are born with scales but shed them as they grow. Filefish have rough non-overlapping scales with small spikes, which is why they are called filefish. Some filefish appear scaleless because their scales are so small. Prominent scaling appears on
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
only along the lateral line and in the
corselet In women's clothing, a corselet or corselette is a type of foundation garment, sharing elements of both bras and girdles. It extends from straps over the shoulders down the torso, and stops around the top of the legs. It may incorporate lace in ...
, a protective band of thickened and enlarged scales in the shoulder region. Over most of their body tuna have scales so small that to casual inspection they seems scaleless. File:Cantherhines fronticinctus.jpg, Some
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 rel ...
appear scaleless because their scales are so small. File:Bluefin-big.jpg, To casual examination
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
seem largely free of scales, but they are not.


Leviticus

A passage in Leviticus declares that, "of all that are in the waters... in the seas, and in the rivers" those that do not have both fins and scales "shall be an abomination unto you" and may not be eaten. This eliminates all aquatic
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s as abominations and unclean, as well as any fish that lack scales (there do not seem to be fish that lack fins). According to the ''chok'' or divine decrees of the
Torah The Torah (; hbo, ''Tōrā'', "Instruction", "Teaching" or "Law") is the compilation of the first five books of the Hebrew Bible, namely the books of Genesis, Exodus, Leviticus, Numbers and Deuteronomy. In that sense, Torah means the s ...
and the
Talmud The Talmud (; he, , Talmūḏ) is the central text of Rabbinic Judaism and the primary source of Jewish religious law (''halakha'') and Jewish theology. Until the advent of modernity, in nearly all Jewish communities, the Talmud was the cente ...
, for a fish to be declared
kosher (also or , ) is a set of dietary laws dealing with the foods that Jewish people are permitted to eat and how those foods must be prepared according to Jewish law. Food that may be consumed is deemed kosher ( in English, yi, כּשר), fro ...
, it must have scales and fins.Aryeh Citron
"All About Kosher Fish"
/ref> The definition of "scale" differs from the definitions presented in biology, in that the scales of a kosher fish must be visible to the eye, and can be easily removed from the skin either by hand or scaling knife. According to the
kosher certification agency A kosher certification agency is an organization or certifying authority that grants a ''hechsher'' ( he, הכשר, "seal of approval") to ingredients, packaged foods, beverages, and certain materials, as well as food-service providers and faciliti ...
of the
Orthodox Union The Orthodox Union (abbreviated OU) is one of the largest Orthodox Jewish organizations in the United States. Founded in 1898, the OU supports a network of synagogues, youth programs, Jewish and Religious Zionist advocacy programs, programs for ...
, a fish is kosher if the scales can be removed without tearing its skin. Thus carp and salmon are kosher, whereas a shark, whose scales are microscopic, a sturgeon, whose scutes can not be easily removed without cutting them out of the body, are all not kosher. Other non-kosher fish include catfish, eels, freshwater cod, snake mackerels, and puffer fish.


Lepidophagy

Lepidophagy Lepidophagy is a specialised feeding behaviour in fish that involves eating the scales of other fish. Lepidophagy is widespread, having evolved independently in at least five freshwater families and seven marine families. A related feeding behav ...
(Ancient Greek for ''scale-eating'') is a specialised feeding behaviour in fish that involves eating the scales of other fish. Lepidophagy has independently
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
in at least five freshwater families and seven marine families. Fish scales can be nutritious, containing a dermal portion and a layer of protein-rich mucus apart from the layers of
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 scales, hair, nails, feathers, ho ...
and enamel. They are a rich source of
calcium phosphate The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are white ...
. However, the energy expended to make a strike versus the amount of scales consumed per strike puts a limit on the size of lepidophagous fish, and they are usually much smaller than their prey. Scale eating behaviour usually evolves because of lack of food and extreme environmental conditions. The eating of scales and the skin surrounding the scales provides protein rich nutrients that may not be available elsewhere in the niche.
Fish jaw Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by b ...
s normally show
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
. An exception occurs with the scale-eating
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
''
Perissodus microlepis ''Perissodus microlepis'' is a species of cichlid endemic to Lake Tanganyika. This species reaches a length of TL. This species can also be found in the aquarium trade. It is a scale-eating 'parasite' on other fish species. It occurs in two ...
''. The jaws of this fish occur in two distinct morphological forms. One morph has its jaw twisted to the left, allowing it to eat scales more readily on its victim's right flank. The other morph has its jaw twisted to the right, which makes it easier to eat scales on its victim's left flank. The relative abundance of the two morphs in populations is regulated by
frequency-dependent selection Frequency-dependent selection is an evolutionary process by which the fitness (biology), fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population. * In positive frequency-dependent selection, the fit ...
.


See also

* Age determination in fish *
Animal coloration Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male h ...
*
Animal reflectors Animal reflectors or mirrors are important to the survival of many kinds of animal, and, in some cases, have been mimicked by engineers developing photonic crystals. Examples are the scales of silvery fish, and the tapetum lucidum that causes the ey ...
*
Photonic crystals A photonic crystal is an optical nanostructure in which the refractive index changes periodically. This affects the propagation of light in the same way that the structure of natural crystals gives rise to X-ray diffraction and that the atomi ...
*
Reptile scale Reptile skin is covered with scutes or scales which, along with many other characteristics, distinguish reptiles from animals of other classes. They are made of alpha and beta-keratin and are formed from the epidermis (contrary to fish, in which t ...
*
Scale (zoology) In most biological nomenclature, a scale ( grc, λεπίς, lepís; la, squāma) is a small rigid plate that grows out of an animal's skin to provide protection. In lepidopteran (butterfly and moth) species, scales are plates on the surface ...
*
Scale armour Scale armour (or scale mail) is an early form of armour consisting of many individual small armour scales (plates) of various shapes attached to each other and to a backing of cloth or leather in overlapping rows.Snake scales Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joints ...
* Urokotori – Japanese fish scaler


References


Further reading

* *


External links

* Hydrodynamic aspects of shark scale

*Fish scales and flow manipulatio

{{Authority control Fish anatomy