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Chromatophore

From Wikipedia, the free encyclopedia
Cells with a primary function of coloration found in a wide range of animals
This article is about a type of cell or multicellular organ. For other uses, seeChromatophore (disambiguation).
Chromatophores in the skin of asquid

Chromatophores are cells that produce color, of which many types arepigment-containing cells, or groups of cells, found in a wide range of animals includingamphibians,fish,reptiles,crustaceans andcephalopods.Mammals andbirds, in contrast, have a class of cells calledmelanocytes forcoloration.

Chromatophores are largely responsible for generating skin andeye colour inectothermic animals and are generated in theneural crest duringembryonic development. Mature chromatophores are grouped into subclasses based on their colour under white light:xanthophores (yellow),erythrophores (red),iridophores (reflective /iridescent),leucophores (white),melanophores (black/brown), andcyanophores (blue). While most chromatophores contain pigments that absorb specific wavelengths of light, the color of leucophores and iridophores is produced by their respective scattering and optical interference properties.

7x speed timelapse video of fish melanophores responding to 200μM adrenaline

Some species can rapidly change colour through mechanisms that translocate pigment and reorient reflective plates within chromatophores. This process, often used as a type ofcamouflage, is calledphysiological colour change ormetachrosis.[1] Cephalopods, such as theoctopus, have complex chromatophore organs controlled by muscles to achieve this, whereas vertebrates such aschameleons generate a similar effect bycell signalling. Such signals can behormones orneurotransmitters and may be initiated by changes in mood, temperature, stress or visible changes in the local environment.[citation needed] Chromatophores are studied by scientists to understand human disease and as a tool indrug discovery.

Human discovery

[edit]

Aristotle mentioned the ability of theoctopus to change colour for bothcamouflage and signalling in hisHistoria animalium (ca 4th century BC):[2]

The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed.

Giosuè Sangiovanni was the first to describeinvertebrate pigment-bearing cells ascromoforo in an Italian science journal in 1819.[3]

Charles Darwin described the colour-changing abilities of thecuttlefish inThe Voyage of the Beagle (1860):[4]

These animals also escape detection by a very extraordinary, chameleon-like power of changing their colour. They appear to vary their tints according to the nature of the ground over which they pass: when in deep water, their general shade was brownish purple, but when placed on the land, or in shallow water, this dark tint changed into one of a yellowish green. The colour, examined more carefully, was a French grey, with numerous minute spots of bright yellow: the former of these varied in intensity; the latter entirely disappeared and appeared again by turns. These changes were effected in such a manner that clouds, varying in tint between a hyacinth red and a chestnut-brown, were continually passing over the body. Any part, being subjected to a slight shock of galvanism, became almost black: a similar effect, but in a less degree, was produced by scratching the skin with a needle. These clouds, or blushes as they may be called, are said to be produced by the alternate expansion and contraction of minute vesicles containing variously coloured fluids.

Classification of chromatophore

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Aveiled chameleon,Chamaeleo calyptratus. Structural green and blue colours are generated by overlaying chromatophore types to reflect filtered light.

The termchromatophore was adopted (following Sangiovanni'schromoforo) as the name for pigment-bearing cells derived from the neural crest of cold-bloodedvertebrates and cephalopods. The word itself comes from theGreek wordschrōma (χρῶμα) meaning "colour," andphoros (φόρος) meaning "bearing". In contrast, the wordchromatocyte (kytos (κύτος) meaning "cell") was adopted for the cells responsible for colour found in birds and mammals. Only one such cell type, themelanocyte, has been identified in these animals.

It was only in the 1960s that chromatophores were well enough understood to enable them to be classified based on their appearance. This classification system persists to this day, even though thebiochemistry of the pigments may be more useful to a scientific understanding of how the cells function.[5]

Colour-producing molecules fall into two distinct classes:biochromes andstructural colours or "schemochromes".[6] The biochromes include true pigments, such ascarotenoids andpteridines. These pigments selectively absorb parts of thevisible light spectrum that makes up white light while permitting otherwavelengths to reach the eye of the observer. Structural colours are produced by various combinations of diffraction, reflection or scattering of light from structures with a scale around a quarter of the wavelength of light. Many such structures interfere with some wavelengths (colours) of light and transmit others, simply because of their scale, so they often produceiridescence by creating different colours when seen from different directions.[citation needed]

Whereas all chromatophores contain pigments or reflecting structures (except when there has been amutation, as inalbinism), not all pigment-containing cells are chromatophores.Haem, for example, is a biochrome responsible for the red appearance of blood. It is found primarily inred blood cells (erythrocytes), which are generated in bone marrow throughout the life of an organism, rather than being formed during embryological development. Therefore, erythrocytes are not classified as chromatophores.[citation needed]

Xanthophores and erythrophores

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Chromatophores that contain large amounts ofyellow pteridine pigments are named xanthophores; those with mainlyred/orange carotenoids are termed erythrophores.[5] However,vesicles containing pteridine and carotenoids are sometimes found in the same cell, in which case the overall colour depends on the ratio of red and yellow pigments.[7] Therefore, the distinction between these chromatophore types is not always clear.

Most chromatophores can generate pteridines fromguanosine triphosphate, but xanthophores appear to have supplemental biochemical pathways enabling them to accumulate yellow pigment. In contrast, carotenoids aremetabolised and transported to erythrophores. This was first demonstrated by rearing normally green frogs on a diet ofcarotene-restrictedcrickets. The absence of carotene in the frogs' diet meant that the red/orange carotenoid colour 'filter' was not present in their erythrophores. This made the frogs appear blue instead of green.[8]

Iridophores and leucophores

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Leucophore layer composition

Iridophores, sometimes also called guanophores, are chromatophores that reflect light using plates of crystalline chemochromes made fromguanine.[9] When illuminated they generate iridescent colours because of the constructive interference of light. Fish iridophores are typically stacked guanine plates separated by layers of cytoplasm to form microscopic, one-dimensional,Bragg mirrors. Both the orientation and the optical thickness of the chemochrome determines the nature of the colour observed.[10] By using biochromes as coloured filters, iridophores create an optical effect known asTyndall orRayleigh scattering, producing bright-blue or -green colours.[11]

A related type of chromatophore, the leucophore, is found in some fish, in particular in thetapetum lucidum. Like iridophores, they utilize crystallinepurines (often guanine) to reflect light. Unlike iridophores, leucophores have more organized crystals that reduce diffraction. Given a source of white light, they produce awhite shine. As with xanthophores and erythrophores, in fish the distinction between iridophores and leucophores is not always obvious, but, in general, iridophores are considered to generate iridescent ormetallic colours, whereas leucophores produce reflective white hues.[11]

Melanophores

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At the bottom amutant zebrafish larva that fails to synthesise melanin in its melanophores, at the top a non-mutant, wildtype larva
See also:Melanocyte

Melanophores containeumelanin, a type ofmelanin, that appearsblack or dark-brown because of its light absorbing qualities. It is packaged in vesicles called melanosomes and distributed throughout the cell. Eumelanin is generated fromtyrosine in a series of catalysed chemical reactions. It is a complex chemical containing units ofdihydroxyindole and dihydroxyindole-2-carboxylic acid with somepyrrole rings.[12] The key enzyme in melanin synthesis istyrosinase. When this protein is defective, no melanin can be generated resulting in certain types of albinism. In some amphibian species there are other pigments packaged alongside eumelanin. For example, a novel deep (wine) red-colour pigment was identified in the melanophores ofphyllomedusine frogs.[13] Some species of anole lizards, such as theAnolis grahami, use melanocytes in response to certain signals and hormonal changes, and is capable of becoming colors ranging from bright blue, brown, and black. This was subsequently identified aspterorhodin, a pteridinedimer that accumulates around eumelanin core, and it is also present in a variety oftree frog species fromAustralia andPapua New Guinea. While it is likely that other lesser-studied species have complex melanophore pigments, it is nevertheless true that the majority of melanophores studied to date do contain eumelanin exclusively.[14]

Humans have only one class of pigment cell, the mammalian equivalent of melanophores, to generate skin, hair, and eye colour. For this reason, and because the large number and contrasting colour of the cells usually make them very easy to visualise, melanophores are by far the most widely studied chromatophore. However, there are differences between the biology of melanophores and that ofmelanocytes. In addition to eumelanin, melanocytes can generate a yellow/red pigment calledphaeomelanin.[citation needed]

The purple-striped dottyback,Pseudochromis diadema, generates itsviolet stripe with an unusual type of chromatophore.

Cyanophores

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Nearly all the vibrant blues in animals and plants are created bystructural coloration rather than by pigments. However, some types ofSynchiropus splendidus do possess vesicles of acyan biochrome of unknown chemical structure in cells named cyanophores.[11] Although they appear unusual in their limited taxonomic range, there may be cyanophores (as well as further unusual chromatophore types) in other fish and amphibians. For example, brightly coloured chromatophores with undefined pigments are found in bothpoison dart frogs andglass frogs,[15] and atypicaldichromatic chromatophores, namederythro-iridophores have been described inPseudochromis diadema.[16]

Pigment translocation

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Fish and frog melanophores are cells that can change colour by dispersing or aggregating pigment-containing bodies.

Many species are able to translocate the pigment inside their chromatophores, resulting in an apparent change in body colour. This process, known asphysiological colour change, is most widely studied in melanophores, since melanin is the darkest and most visible pigment. In most species with a relatively thindermis, the dermal melanophores tend to be flat and cover a large surface area. However, in animals with thick dermal layers, such as adult reptiles, dermal melanophores often form three-dimensional units with other chromatophores. These dermal chromatophore units (DCU) consist of an uppermost xanthophore or erythrophore layer, then an iridophore layer, and finally a basket-like melanophore layer with processes covering the iridophores.[17]

Both types of melanophore are important in physiological colour change. Flat dermal melanophores often overlay other chromatophores, so when the pigment is dispersed throughout the cell the skin appears dark. When the pigment is aggregated toward the centre of the cell, the pigments in other chromatophores are exposed to light and the skin takes on their hue. Likewise, after melanin aggregation in DCUs, the skin appears green through xanthophore (yellow) filtering of scattered light from the iridophore layer. On the dispersion of melanin, the light is no longer scattered and the skin appears dark. As the other biochromatic chromatophores are also capable of pigment translocation, animals with multiple chromatophore types can generate a spectacular array of skin colours by making good use of the divisional effect.[18][19]

A singlezebrafish melanophore imaged bytime-lapse photography during pigment aggregation

The control and mechanics of rapid pigment translocation has been well studied in a number of different species, in particular amphibians andteleost fish.[11][20] It has been demonstrated that the process can be underhormonal orneuronal control or both and for many species of bony fishes it is known that chromatophores can respond directly to environmental stimuli like visible light, UV-radiation, temperature, pH, chemicals, etc.[21] Neurochemicals that are known to translocate pigment includenoradrenaline, through itsreceptor on the surface on melanophores.[22] The primary hormones involved in regulating translocation appear to be themelanocortins,melatonin, andmelanin-concentrating hormone (MCH), that are produced mainly in the pituitary, pineal gland, and hypothalamus, respectively. These hormones may also be generated in aparacrine fashion by cells in the skin. At the surface of the melanophore, the hormones have been shown to activate specificG-protein-coupled receptors that, in turn, transduce the signal into the cell. Melanocortins result in the dispersion of pigment, while melatonin and MCH results in aggregation.[23]

Numerous melanocortin, MCH and melatonin receptors have been identified in fish[24] and frogs,[25] including ahomologue ofMC1R,[26] a melanocortin receptor known to regulateskin andhair colour in humans.[27] It has been demonstrated thatMC1R is required in zebrafish for dispersion of melanin.[28] Inside the cell,cyclic adenosine monophosphate (cAMP) has been shown to be an importantsecond messenger of pigment translocation. Through a mechanism not yet fully understood, cAMP influences other proteins such asprotein kinase A to drivemolecular motors carrying pigment containing vesicles along bothmicrotubules andmicrofilaments.[29][30][31]

Background adaptation

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Zebrafish chromatophores mediatebackground adaptation on exposure to dark (top) and light environments (bottom).
See also:Camouflage

Most fish, reptiles and amphibians undergo a limited physiological colour change in response to a change in environment. This type of camouflage, known asbackground adaptation, most commonly appears as a slight darkening or lightening of skin tone to approximatelymimic the hue of the immediate environment. It has been demonstrated that the background adaptation process is vision-dependent (it appears the animal needs to be able to see the environment to adapt to it),[32] and that melanin translocation in melanophores is the major factor in colour change.[23] Some animals, such as chameleons andanoles, have a highly developed background adaptation response capable of generating a number of different colours very rapidly.[33] They have adapted the capability to change colour in response to temperature, mood, stress levels, and social cues, rather than to simply mimic their environment.

Development

[edit]
Cross-section of a developing vertebratetrunk showing the dorsolateral (red) and ventromedial (blue) routes of chromatoblast migration

During vertebrateembryonic development, chromatophores are one of a number of cell types generated in theneural crest, a paired strip of cells arising at the margins of theneural tube. These cells have the ability to migrate long distances, allowing chromatophores to populate many organs of the body, including the skin, eye, ear, and brain. Fish melanophores and iridophores have been found to contain the smooth muscle regulatory proteins [calponin] andcaldesmon.[34] Leaving the neural crest in waves, chromatophores take either a dorsolateral route through the dermis, entering theectoderm through small holes in thebasal lamina, or a ventromedial route between thesomites and the neural tube. The exception to this is the melanophores of the retinal pigmented epithelium of the eye. These are not derived from the neural crest. Instead, an outpouching of the neural tube generates theoptic cup, which, in turn, forms theretina.[citation needed]

When and howmultipotent chromatophore precursor cells (calledchromatoblasts) develop into their daughter subtypes is an area of ongoing research. It is known in zebrafish embryos, for example, that by 3 days afterfertilization each of the cell classes found in the adult fish—melanophores, xanthophores and iridophores—are already present. Studies using mutant fish have demonstrated thattranscription factors such askit,sox10, andmitf are important in controlling chromatophore differentiation.[35] If these proteins are defective, chromatophores may be regionally or entirely absent, resulting in aleucistic disorder.

Practical applications

[edit]

Chromatophores are sometimes used in applied research. For example, zebrafish larvae are used to study how chromatophores organise and communicate to accurately generate the regular horizontal striped pattern as seen in adult fish.[36] This is seen as a usefulmodel system for understanding patterning in theevolutionary developmental biology field. Chromatophore biology has also been used to model human condition or disease, includingmelanoma and albinism. Recently, the gene responsible for the melanophore-specificgolden zebrafish strain,Slc24a5, was shown to have a human equivalent that strongly correlates withskin colour.[37]

Chromatophores are also used as abiomarker of blindness in cold-blooded species, as animals with certain visual defects fail to background adapt to light environments.[32] Human homologues of receptors that mediate pigment translocation in melanophores are thought to be involved in processes such asappetite suppression andtanning, making them attractive targets fordrugs.[26] Therefore, pharmaceutical companies have developed abiological assay for rapidly identifying potential bioactive compounds using melanophores from theAfrican clawed frog.[38] Other scientists have developed techniques for using melanophores asbiosensors,[39] and for rapid disease detection (based on the discovery thatpertussis toxin blocks pigment aggregation in fish melanophores).[40] Potentialmilitary applications of chromatophore-mediated colour changes have been proposed, mainly as a type ofactive camouflage, which could as incuttlefish make objects nearly invisible.[41][42]

Cephalopod chromatophores

[edit]
An infantcuttlefish, using background adaptation to mimic the local environment

Coleoid cephalopods (including octopuses,squids andcuttlefish) have complex multicellular organs that they use to change colour rapidly, producing a wide variety of bright colours and patterns. Each chromatophore unit is composed of a single chromatophore cell and numerous muscle, nerve,glial, and sheath cells.[43] Inside the chromatophore cell, pigment granules are enclosed in an elastic sac, called the cytoelastic sacculus. To change colour the animal distorts the sacculus form or size by muscular contraction, changing itstranslucency, reflectivity, oropacity. This differs from the mechanism used in fish, amphibians, and reptiles in that the shape of the sacculus is changed, rather than translocating pigment vesicles within the cell. However, a similar effect is achieved. The energy cost of the complete activation of the chromatophore system is very high, being nearly as much as all the energy used by an octopus at rest.[44]

Octopuses and mostcuttlefish[45] can operate chromatophores in complex, undulating chromatic displays, resulting in a variety of rapidly changing colour schemata. The nerves that operate the chromatophores are thought to be positioned in the brain in a pattern isomorphic to that of the chromatophores they each control. This means the pattern of colour change functionally matches the pattern ofneuronal activation. This may explain why, as the neurons are activated in iterative signal cascade, one may observe waves of colour changing.[46] Like chameleons, cephalopods use physiologicalcolour change for social interaction. They are also among the most skilled at camouflage, having the ability to match both the colour distribution and thetexture of their local environment with remarkable accuracy.

See also

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Notes

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  2. ^Aristotle.Historia Animalium. IX, 622a: 2-10. About 400 BC. Cited in Luciana Borrelli,Francesca Gherardi, Graziano Fiorito.A catalogue of body patterning in Cephalopoda. Firenze University Press, 2006.AbstractArchived 2018-02-06 at theWayback MachineGoogle books
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  4. ^Darwin, Charles (1860). "Chapter 1. Habits of a Sea-slug and Cuttle-fish".Journal Of Researches Into The Natural History And Geology Of The Countries Visited During The Voyage Round The World Of H.M.S. 'Beagle' Under The Command Of Captain Fitz Roy, R.N. John Murray, London. p. 7.
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Vision in animals
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Developmental stages:SpawnParalarva (Doratopsis stage) → Juvenile → Subadult → Adult •Egg fossilsProtoconch (embryonic shell)
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