Fish scale

Source: Wikipedia, the free encyclopedia.
For other uses, see Fish scale (disambiguation).

teleost fish (rohu
)

A fish scale is a small rigid plate that grows out of the

jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as well as possible hydrodynamic advantages. The term scale derives from the Old French escale, meaning a shell pod or husk.[1]

Scales vary enormously in size, shape, structure, and extent, ranging from strong and rigid armour plates in fishes such as

boxfishes, to microscopic or absent in fishes such as eels and anglerfishes. The morphology of a scale can be used to identify the species of fish it came from. Scales originated within the jawless ostracoderms
, ancestors to all jawed fishes today. Most
Cartilaginous fishes (sharks and rays) are covered with placoid scales. Some species are covered instead by scutes
, and others have no outer covering on part or all of the skin.

Fish scales are part of the fish's integumentary system, and are produced from the mesoderm layer of the dermis, which distinguishes them from reptile scales.[2][3] The same genes involved in tooth and hair development in mammals 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 or slime which can protect against pathogens such as bacteria, fungi, and viruses, and reduce surface resistance when the fish swims.

Thelodont scales

Left to right: denticles of Paralogania (?), Shielia taiti, Lanarkia horrida

The bony scales of

fossil fish, are well understood. The scales were formed and shed throughout the organisms' lifetimes, and quickly separated after their death.[4]

Bone, a tissue that is both resistant to mechanical damage and relatively prone to fossilization, often preserves internal detail, which allows the

dentine, with a sometimes-ornamented enameloid upper surface and an aspidine base.[5] Its growing base is made of cell-free bone, which sometimes developed anchorage structures to fix it in the side of the fish.[6] 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.[7] 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.[6]

However, using scale morphology alone to distinguish species has some pitfalls. Within each organism, scale shape varies hugely according to body area,[8] 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.[9]

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.[7] 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.[10]

Cosmoid scales

Queensland lungfish

Cosmoid scales are found only on ancient

vitrodentine. The upper surface is keratin. Cosmoid scales increase in size through the growth of the lamellar bone layer.[11]

Elasmoid scales

Lobe-finned fishes, like this preserved coelacanth
, have elasmoid scales.

Elasmoid scales are thin,

lobe-finned fishes, such as all extant and some extinct lungfishes, as well as the coelacanths which have modified cosmoid scales that lack cosmine and are thinner than true cosmoid scales. They are also present in some tetrapodomorphs like Eusthenopteron
, amiids, and teleosts, whose cycloid and ctenoid scales represent the least mineralized elasmoid scales.

The zebrafish elasmoid scales are used in the lab to study bone mineralization process, and can be cultured (kept) outside of the organism.[13][14]

Ganoid scales

The scales of this spotted gar appear glassy due to ganoine.
Mineral texture of ganoine layers in the scales of an alligator gar

Ganoid scales are found in the

vitrodentine
.

Ganoine is a characteristic component of ganoid scales. It is a glassy, often multi-layered mineralized

synapomorphic character of ray-finned fishes, ganoine or ganoine-like tissues are also found on the extinct acanthodii.[17] It has been suggested ganoine is homologous to tooth enamel in vertebrates[15] or even considered a type of enamel.[18]



Amblypterus striatus
 Ganoid scales of the extinct Carboniferous fish, Amblypterus striatus. (a) shows the outer surface of four of the scales, and (b) shows the inner surface of two of the scales. Each of the rhomboidal-shaped ganoid scales of Amblypterus has a ridge on the inner surface which is produced at one end into a projecting peg which fits into a notch in the next scale, similar to the manner in which tiles are pegged together on the roof of a house.

Most ganoid scales are

rhomboidal (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.[19]
In this way, ganoid scales are nearly impenetrable and are excellent protection against predation.

  • The alligator gar has a tough armouring of rhomboidal-shaped ganoid scales.[19]
    The
    rhomboidal-shaped ganoid scales.[19]
  • The sturgeon has rows of ganoid scales enlarged into scute-like armour plates.
    The sturgeon has rows of ganoid scales enlarged into scute-like armour plates.
  • The ganoid scales on a bowfin are reduced in size and resemble cycloid scales.
    The ganoid scales on a
    cycloid scales
    .
Geometrically laid out ganoid scales on a bichir

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.

  • Earrings made from the ganoid scales of an alligator gar
    Earrings made from the ganoid scales of an alligator gar
  • Fossil of a primitive rayfin with ganoid scales
    Fossil of a primitive rayfin with ganoid scales
  • Ganoid scales on a fossilised Lepidotes, circa. 130 mya
    Ganoid scales on a fossilised
    mya

Native Americans and people of the Caribbean used the tough ganoid scales of the alligator gar for arrow heads, breastplates, and as shielding to cover plows. In current times jewellery is made from these scales.[20]

Leptoid scales

Leptoid (bony-ridge) scales are found on higher-order bony fish, the

derived clade 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.[21]

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

age the fish
.

Leptoid scales come in two forms: cycloid (smooth) and ctenoid (comb-like).[23]

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 and carp.

Asian arowana have large cycloid scales arranged on the fish in a mosaic of raised ribs (left). The scales themselves are covered with a delicate net pattern (right).[24][25]
Cycloid (circular) scales
carp-like
fish with circular cycloid scales that are smooth to the touch.
Cycloid (circular) scales are usually found on carp-like or salmon-like fishes.

Ctenoid scales

Ctenoid (toothed) scales are like cycloid scales, except they have small teeth or spinules 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 and calcium carbonate and a deeper layer composed mostly of collagen. The enamel of the other scale types is reduced to superficial ridges and ctenii.

Ctenoid (toothed) scales
The ctenoid scale of a perch has a toothed outer edge (at top of image).
This dottyback is a perch-like fish with toothed ctenoid scales that are rough to the touch.
The size of the teeth on ctenoid scales can vary with position, as these scales from the
rattail
Cetonurus crassiceps show.
Ctenoid scales from a perch vary from the medial (middle of the fish), to dorsal (top), to caudal (tail end) scales.
Crazy fish have cycloid scales on the belly but ctenoid scales elsewhere.[26]
Ctenoid (toothed) scales are usually found on perch-like fishes.
 

Ctenoid scales, similar to other epidermal structures, originate from

mineralization.[27] The circumference of the scales grows first, followed by thickness when overlapping layers mineralize together.[27]

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 flatfishes 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

The herring's reflectors are nearly vertical for camouflage from the side.
The deep sea hatchetfish has scales which reflect blue light.
The scales of a typical teleost fish, like this Atlantic herring, are silvered.

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.[29]

The marine hatchetfish is extremely flattened laterally (side to side), leaving the body just millimetres thick, and the body is so silvery as to resemble aluminium foil. The mirrors consist of microscopic structures similar to those used to provide structural coloration: stacks of between 5 and 10 crystals of guanine 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.[29]

Most fish in the upper ocean are camouflaged by silvering. In fish such as the herring, 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.[29]

Fish scales with these properties are used in some cosmetics, since they can give a shimmering effect to makeup and lipstick.[30]

Placoid scales

Placoid scales as viewed through an electron microscope. Also called dermal denticles, these are structurally homologous with vertebrate teeth.

Placoid (pointed, tooth-shaped) scales are found in the

vitrodentine, 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

placoderms
.

Shark skin

Cartilaginous fishes, like this tiger shark, have placoid scales (dermal denticles).

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

anti-fouling by exhibiting the lotus effect.[32]

All denticles are composed of an interior pulp cavity with a nervous and arterial supply rooted in the dermis to supply the denticle with mucus.[33] 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.[34] 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.[34] Both riblet shapes assist in creating a turbulent boundary layer forcing the laminar flow farther away from the sharks skin.[35]

Unlike bony fish, sharks have a complicated dermal corset made of flexible collagenous fibers arranged as a helical network surrounding their body. The corset works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy.[36] 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.[37]

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.[38]

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.[39] The crown and the neck of the denticles however play a key role and are responsible for creating the turbulent vortices and eddies found near the skin's surface.[39] 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. During a recent research experiment biomimetic samples of shark denticles with a crescent like microstructure were 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.[40] 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%.[41]

Drag reduction

Effects of turbulent flow on boundary layer
Diagram of the side profile of a shark denticle showing a vortex in the wake downstream of the denticle

Sharks decrease drag and overall

Pressure drag is created from the pressure difference between the anterior and posterior sides of the shark due to the amount of volume that is pushed past the shark to propel itself forward.[42] This type of drag is also directly proportional to the laminar flow. When the laminar flow increases around the fish the pressure drag does as well.[43] 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.[42]

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.[38]

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 are abruptly liquidated 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.[44]

Technical application

The rough, sandpaper-like texture of shark and ray skin, coupled with its toughness, has led it to be valued as a source of rawhide leather, called shagreen. One of the many historical applications of shark shagreen was in making hand-grips for swords. The rough texture of the skin is also used in Japanese cuisine to make graters called oroshiki, by attaching pieces of shark skin to wooden boards. The small size of the scales grates the food very finely.

Barnacle growth on boat hull

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 barnacles, algae, 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 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%.[45]

There are many examples of biomimetic materials 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 from coating the hulls of submarines and ships. One variety is traded as "sharklet".[46][47]

A lot of the new methods for replicating shark skin involve the use of polydimethylsiloxane (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 properties, exhibiting the lotus effect.[46] One study found that these biomimetic surfaces reduced drag by up to 9%,[37] while with flapping motion drag reduction reached 12.3%.[48]

Denticles also provide drag reduction on objects where the main form of drag is caused by turbulent flow at the surface. A large portion of the total drag on long objects with relatively flat sides usually comes from turbulence at the wall, so riblets will have an appreciable effect. Along with marine applications, the aerospace industry can benefit greatly from these biomimetic designs. Other applications include pipes, where they score the insides to a riblet-like roughness and have discovered a 5% drag reduction, and a few percent reduction is claimed with competitive swimwear.[49]

Parametric modeling has been done on shark denticles with a wide range of design variations such as low and high-profile vortex generators.[50] Through this method, the most thorough characterization has been completed for symmetrical two-dimensional riblets with sawtooth, scalloped and blade cross sections.[51]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 reacted. Both the geometry of the denticles and their arrangement have a profound effect on the aerodynamic response of the aerofoils. 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 denticle's wake and stream-wise vortices that replenish momentum lost in the boundary layer due to skin friction.[50]

Scutes

Pineconefish
are covered in scutes.

Scutes 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, are completely or partially covered in scutes. River herrings and threadfins 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
on either side.

Scale development

Scales typically appear late in the development of fish. In the case of zebrafish, 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 occurs, then morphogenesis is induced, and finally the process of differentiation or late metamorphosis occurs.[52][53]

  • Mesenchyme consolidation: The consolidation or structuring of the mesenchyme originates during the development of the
    collagen fibers. Subsequently, for both fish the fibroblasts 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.[52][53][54]
  • Morphogenesis induction: The morphogenesis is due to the formation of the epidermal papilla, which is generated by joining the epidermis and dermis through a process of invagination. 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 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 will arise.[52][53][54]
  • 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.[52][53][54]

Unlike elasmoid scales,

Sharpey fibers.[52][53][54]

One of the genes that regulate the development of scale formation in fish is the

cell differentiation and interaction, the level of ApoE transcription is high, which has led to the conclusion that this protein is important for the late development of scales.[55][56]

Modified scales

Lateral line
Cycloid scales of a common roach. Modified scales along the lateral line are visible in the lower half.
Closeup of a modified cycloid scale from the lateral line of a wrasse
Surgeonfish
Surgeonfish (left) have a sharp, scalpel-like modified scale on either side just before the tail. Closeup (right).

Different groups of fish have

evolved
a number of modified scales to serve various functions.

  • Porcupine fish have scales modified into spines.
    Porcupine fish have scales modified into spines
    .
  • Pufferfish spines are not modified scales but are developed by an independent gene network.
    Pufferfish
    spines are not modified scales but are developed by an independent gene network.

Fish without scales

  • Mandarinfish lack scales and protect themselves with a layer of smelly and bitter slime.
    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 (lampreys and hagfishes) have smooth skin without scales and without dermal bone.[60] Lampreys get some protection from a tough leathery skin. Hagfish exude copious quantities slime or mucus if they are threatened.[61] They can tie themselves in an overhand knot, scraping off the slime as they go and freeing themselves from a predator.[62]
  • Most eels are scaleless, though some species are covered with tiny smooth cycloid scales.
  • Most catfish lack scales, though several families have body armour in the form of dermal plates or some sort of scute.[63]
  • aposematic.[64]
  • Anglerfish have loose, thin skin often covered with fine forked dermal prickles or tubercles, 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

poachers, and several families of sticklebacks
, have developed external bony plates, structurally resembling placoid scales, as protective armour against predators.

Eels seem scaleless, but some species are covered with tiny smooth cycloid scales.
  • Boxfish have plates of ossified skin fused together to form a rigid shell.
    Boxfish
    have plates of ossified skin fused together to form a rigid shell.
  • Seahorses have thin skin stretched over bony plates arranged in rings.
    Seahorses 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 only along the lateral line and in the corselet, 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.[65]

  • Some filefish appear scaleless because their scales are so small.
    Some filefish appear scaleless because their scales are so small.
  • To casual examination tuna seem largely free of scales, but they are not.
    To casual examination tuna 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.[66] This eliminates all aquatic invertebrates 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

kosher, it must have scales and fins.[67] 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.[67] According to the kosher certification agency of the Orthodox Union, a fish is kosher if the scales can be removed without tearing its skin.[68] 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.[67]

Lepidophagy

Dorsal view of right-bending (left) and left-bending (right) jaw morphs adapted for eating fish scales[69]

evolved in at least five freshwater families and seven marine families.[71]

Fish scales can be nutritious, containing a dermal portion and a layer of protein-rich mucus apart from the layers of

much smaller than their prey.[71] 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.[72]

bilateral symmetry. An exception occurs with the scale-eating cichlid Perissodus microlepis. 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.[69][73][74]

See also

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Further reading