Biological pigments, also known simply as pigments or biochromes, are substances produced by living organisms that have a

color Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associ ...

resulting from selective color absorption. Biological pigments include plant pigments and flower pigments. Many biological structures, such as

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 ...

eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...

feathers Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier e ...

fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...

and

hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...

contain pigments such as

melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...

in specialized cells called

chromatophore Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...

s. In some species, pigments accrue over very long periods during an individual's lifespan. Pigment color differs from

structural color 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 ...

in that it is the same for all viewing angles, whereas structural color is the result of selective

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 s ...

iridescence Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...

, usually because of multilayer structures. For example,

butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...

wings typically contain structural color, although many butterflies have cells that contain pigment as well.

Biological pigments

See

conjugated system In theoretical chemistry, a conjugated system is a system of connected p-orbitals with delocalized electrons in a molecule, which in general lowers the overall energy of the molecule and increases stability. It is conventionally represented as ...

s for electron bond chemistry that causes these molecules to have pigment. *

Heme Heme, or haem (pronounced / hi:m/ ), is a precursor to hemoglobin, which is necessary to bind oxygen in the bloodstream. Heme is biosynthesized in both the bone marrow and the liver. In biochemical terms, heme is a coordination complex "consisti ...

porphyrin Porphyrins ( ) are a group of heterocyclic macrocycle organic compounds, composed of four modified pyrrole subunits interconnected at their α carbon atoms via methine bridges (=CH−). The parent of porphyrin is porphine, a rare chemical com ...

-based:

chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...

bilirubin Bilirubin (BR) (Latin for "red bile") is a red-orange compound that occurs in the normal catabolic pathway that breaks down heme in vertebrates. This catabolism is a necessary process in the body's clearance of waste products that arise from the ...

hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...

hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...

myoglobin Myoglobin (symbol Mb or MB) is an iron- and oxygen-binding protein found in the cardiac and skeletal muscle tissue of vertebrates in general and in almost all mammals. Myoglobin is distantly related to hemoglobin. Compared to hemoglobin, myoglobi ...

* Light-emitting:

luciferin Luciferin (from the Latin ''lucifer'', "light-bearer") is a generic term for the light-emitting compound found in organisms that generate bioluminescence. Luciferins typically undergo an enzyme-catalyzed reaction with molecular oxygen. The result ...

Carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, and Fungus, fungi. Carotenoids give the characteristic color to pumpki ...

s: **

Hematochrome Hematochrome is a yellow, orange, or (most commonly) red biological pigment present in some green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta ...

s (algal pigments, mixes of carotenoids and their derivates) **

Carotene The term carotene (also carotin, from the Latin ''carota'', "carrot") is used for many related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but in general cannot be made by animals (with the exc ...

s: alpha and beta

carotene The term carotene (also carotin, from the Latin ''carota'', "carrot") is used for many related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but in general cannot be made by animals (with the exc ...

lycopene Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene (from the neo-Latin ''Lycopersicum'', the tomato species) is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and vegetables. Occu ...

rhodopsin Rhodopsin, also known as visual purple, is a protein encoded by the RHO gene and a G-protein-coupled receptor (GPCR). It is the opsin of the rod cells in the retina and a light-sensitive receptor protein that triggers visual phototransduction ...

Xanthophyll Xanthophylls (originally phylloxanthins) are yellow pigments that occur widely in nature and form one of two major divisions of the carotenoid group; the other division is formed by the carotenes. The name is from Greek (, "yellow") and (, "l ...

canthaxanthin Canthaxanthin is a keto-carotenoid pigment widely distributed in nature. Carotenoids belong to a larger class of phytochemicals known as terpenoids. The chemical formula of canthaxanthin is C40H52O2. It was first isolated in edible mushrooms. It ...

zeaxanthin Zeaxanthin is one of the most common carotenoids in nature, and is used in the xanthophyll cycle. Synthesized in plants and some micro-organisms, it is the pigment that gives paprika (made from bell peppers), corn, saffron, goji ( wolfberries), ...

lutein Lutein (;"Lutein"

phytochrome Phytochromes are a class of photoreceptor in plants, bacteria and fungi used to detect light. They are sensitive to light in the red and far-red region of the visible spectrum and can be classed as either Type I, which are activated by far-re ...

phycobiliprotein Phycobiliproteins are water-soluble proteins present in cyanobacteria and certain algae (rhodophytes, cryptomonads, glaucocystophytes). They capture light energy, which is then passed on to chlorophylls during photosynthesis. Phycobiliproteins are ...

s *

Psittacofulvin Psittacofulvin pigments, sometimes called psittacins are responsible for the bright-red, orange, and yellow colours specific to parrots. In parrots, psittacofulvins are synthesized by a polyketide synthase enzyme that is expressed in growing feath ...

s: a class of red and yellow pigments unique to

parrot Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genera comprising the order Psittaciformes (), found mostly in tropical and subtropical regions. The order is subdivided into three superfamilies: the Psittacoid ...

s *

Turacin Turacin is a naturally occurring red pigment that is 6% copper complexed to uroporphyrin III. Arthur Herbert Church discovered turacin in 1869. It is found only in the bird family Musophagidae, the turacos. Other birds derive their red coloration ...

and

Turacoverdin Turacoverdin is a unique copper uroporphyrin pigment responsible for the bright green coloration of several birds of the family Musophagidae, most notably the turaco. It is chemically related to turacin, a red pigment also found almost exclusively ...

: red and green pigments found in

turaco The turacos make up the bird family Musophagidae ( "banana-eaters"), which includes plantain-eaters and go-away-birds. In southern Africa both turacos and go-away-birds are commonly known as loeries. They are semi-zygodactylous: the fourth ( ...

s and related species * Other:

urochrome Urobilin or urochrome is the chemical primarily responsible for the yellow color of urine. It is a linear pyrrole, tetrapyrrole compound that, along with the related colorless compound urobilinogen, are Heme#Degradation, degradation products of th ...

flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...

Pigments in plants

The primary function of pigments in plants is

photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...

, which uses the green pigment

and several colorful pigments that absorb as much light energy as possible. Pigments are also known to play a role in pollination where pigment accumulation or loss can lead to

floral color change Floral color change occurs in flowers in a wide range of angiosperm taxa that undergo a color change associated with their age, or after successful pollination. History The first written record of the term ''floral color change'' was in 1877 wh ...

, signaling to pollinators which flowers are rewarding and contain more pollen and nectar. Plant pigments include many molecules, such as

carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, and Fungus, fungi. Carotenoids give the characteristic color to pumpki ...

anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compo ...

s and

betalain Betalains are a class of red and yellow tyrosine-derived pigments found in plants of the order Caryophyllales, where they replace anthocyanin pigments. Betalains also occur in some higher order fungi. They are most often noticeable in the petals o ...

s. All biological pigments selectively absorb certain wavelengths of

light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 tera ...

while reflecting others. The principal pigments responsible are: *

Chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...

is the primary pigment in plants; it is a

chlorin In organic chemistry, chlorins are tetrapyrrole pigments that are partially hydrogenated porphyrins. The parent chlorin is an unstable compound which undergoes air oxidation to porphine. The name chlorin derives from chlorophyll. Chlorophylls a ...

that absorbs blue and red wavelengths of light while reflecting a majority of

green Green is the color between cyan and yellow on the visible spectrum. It is evoked by light which has a dominant wavelength of roughly 495570 Nanometre, nm. In subtractive color systems, used in painting and color printing, it is created by ...

. It is the presence and relative abundance of chlorophyll that gives plants their green color. All land plants and

green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...

possess two forms of this pigment: chlorophyll ''a'' and chlorophyll ''b''.

Kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "underwat ...

diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...

s, and other photosynthetic

heterokont Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which ...

s contain chlorophyll ''c'' instead of ''b'', while red algae possess only chlorophyll ''a''. All chlorophylls serve as the primary means plants use to intercept light in order to fuel photosynthesis. *

s are red, orange, or yellow

tetraterpenoid Tetraterpenes are terpenes consisting of eight isoprene units and have the molecular formula C40H64. Tetraterpenoids (including many carotenoids) are tetraterpenes that have been chemically modified, as indicated by the presence of oxygen-contain ...

s. During the process of photosynthesis, they have functions in light-harvesting (as

accessory pigments Accessory pigments are light-absorbing compounds, found in photosynthetic organisms, that work in conjunction with chlorophyll ''a''. They include other forms of this pigment, such as chlorophyll ''b'' in green algal and higher plant antennae, w ...

), in

photoprotection Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative stre ...

(energy dissipation via

non-photochemical quenching Non-photochemical quenching (NPQ) is a mechanism employed by plants and algae to protect themselves from the adverse effects of high light intensity. It involves the quenching of singlet excited state chlorophylls (Chl) via enhanced internal con ...

as well as singlet oxygen scavenging for prevention of photooxidative damage), and also serve as protein structural elements. In higher plants, they also serve as precursors to the plant hormone

abscisic acid Abscisic acid (ABA) is a plant hormone. ABA functions in many plant developmental processes, including seed and bud dormancy, the control of organ size and stomatal closure. It is especially important for plants in the response to environmental s ...

. *

Betalain Betalains are a class of red and yellow tyrosine-derived pigments found in plants of the order Caryophyllales, where they replace anthocyanin pigments. Betalains also occur in some higher order fungi. They are most often noticeable in the petals o ...

s are red or yellow pigments. Like anthocyanins they are water-soluble, but unlike anthocyanins they are synthesized from

tyrosine -Tyrosine or tyrosine (symbol Tyr or Y) or 4-hydroxyphenylalanine is one of the 20 standard amino acids that are used by cells to synthesize proteins. It is a non-essential amino acid with a polar side group. The word "tyrosine" is from the Gr ...

. This class of pigments is found only in the

Caryophyllales Caryophyllales ( ) is a diverse and heterogeneous order of flowering plants that includes the cacti, carnations, amaranths, ice plants, beets, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves. The betalai ...

(including

cactus A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Greek ...

and

amaranth ''Amaranthus'' is a cosmopolitan genus of annual or short-lived perennial plants collectively known as amaranths. Some amaranth species are cultivated as leaf vegetables, pseudocereals, and ornamental plants. Catkin-like cymes of densely pack ...

), and never co-occur in plants with anthocyanins. Betalains are responsible for the deep red color of

beet The beetroot is the taproot portion of a beet plant, usually known in North America as beets while the vegetable is referred to as beetroot in British English, and also known as the table beet, garden beet, red beet, dinner beet or golden beet ...

s. *

Anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compo ...

s (literally "flower blue") are

water-soluble In chemistry, solubility is the ability of a substance, the solute, to form a solution with another substance, the solvent. Insolubility is the opposite property, the inability of the solute to form such a solution. The extent of the solub ...

pigments A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...

that appear red to blue, according to pH. They occur in all tissues of higher plants, providing color in leaves, plant stem, roots, flowers, and fruits, though not always in sufficient quantities to be noticeable. Anthocyanins are most visible in the

petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...

s of flowers of many species. Plants, in general, contain six ubiquitous carotenoids:

neoxanthin Neoxanthin is a carotenoid and xanthophyll. In plants, it is an intermediate in the biosynthesis of the plant hormone abscisic acid. It is often present in two forms: all-trans and 9-cis isomers. It is produced from violaxanthin, but a suspect ...

violaxanthin Violaxanthin is a xanthophyll pigment with an orange color found in a variety of plants. Violaxanthin is the product of the epoxidation of zeaxanthin where the oxygen atoms are from reactive oxygen species (ROS). Such ROS's arise when a plant ...

antheraxanthin Antheraxanthin (from ''ánthos'', Greek for "flower" and ''xanthos'', Greek for "yellow") is a bright yellow accessory pigment found in many organisms that perform photosynthesis. It is a xanthophyll cycle pigment, an oil-soluble alcohol within th ...

lutein Lutein (;"Lutein"

β-carotene β-Carotene is an organic, strongly coloured red-orange pigment abundant in fungi, plants, and fruits. It is a member of the carotenes, which are terpenoids (isoprenoids), synthesized biochemically from eight isoprene units and thus having 40 ...

. Lutein is a yellow pigment found in fruits and vegetables and is the most abundant carotenoid in plants.

Lycopene Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene (from the neo-Latin ''Lycopersicum'', the tomato species) is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and vegetables. Occu ...

is the red pigment responsible for the color of

tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...

es. Other less common carotenoids in plants include lutein epoxide (in many woody species), lactucaxanthin (found in lettuce), and alpha carotene (found in carrots). Bougainvillea-3colors

A particularly noticeable manifestation of pigmentation in plants is seen with

autumn leaf color Autumn leaf color is a phenomenon that affects the normal green leaves of many deciduous trees and shrubs by which they take on, during a few weeks in the autumn season, various shades of yellow, orange, red, purple, and brown. The phenomenon i ...

, a phenomenon that affects the normally

leaves of many

deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

s and

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

s whereby they take on, during a few weeks in the

autumn Autumn, also known as fall in American English and Canadian English, is one of the four temperate seasons on Earth. Outside the tropics, autumn marks the transition from summer to winter, in September ( Northern Hemisphere) or March ( Sou ...

season, various shades of

red Red is the color at the long wavelength end of the visible spectrum of light, next to orange and opposite violet. It has a dominant wavelength of approximately 625–740 nanometres. It is a primary color in the RGB color model and a secondar ...

yellow Yellow is the color between green and orange on the spectrum of light. It is evoked by light with a dominant wavelength of roughly 575585 nm. It is a primary color in subtractive color systems, used in painting or color printing. In the R ...

purple Purple is any of a variety of colors with hue between red and blue. In the RGB color model used in computer and television screens, purples are produced by mixing red and blue light. In the RYB color model historically used by painters, pu ...

, and

brown Brown is a color. It can be considered a composite color, but it is mainly a darker shade of orange. In the CMYK color model used in printing or painting, brown is usually made by combining the colors orange and black. In the RGB color model used ...

. Chlorophylls degrade into colorless

tetrapyrrole Tetrapyrroles are a class of chemical compounds that contain four pyrrole or pyrrole-like rings. The pyrrole/pyrrole derivatives are linked by ( =- or -- units), in either a linear or a cyclic fashion. Pyrroles are a five-atom ring with four car ...

s known as ''nonfluorescent chlorophyll catabolites'' (NCCs). As the predominant chlorophylls degrade, the hidden pigments of yellow

xanthophyll Xanthophylls (originally phylloxanthins) are yellow pigments that occur widely in nature and form one of two major divisions of the carotenoid group; the other division is formed by the carotenes. The name is from Greek (, "yellow") and (, "l ...

s and orange beta-carotene are revealed. These pigments are present throughout the year, but the red pigments, the

s, are synthesized ''de novo'' once roughly half of chlorophyll has been degraded. The

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

s released from degradation of light harvesting complexes are stored all winter in the tree's roots, branches, stems, and

trunk Trunk may refer to: Biology * Trunk (anatomy), synonym for torso * Trunk (botany), a tree's central superstructure * Trunk of corpus callosum, in neuroanatomy * Elephant trunk, the proboscis of an elephant Computing * Trunk (software), in rev ...

until next spring when they are recycled to re‑leaf the tree.

Pigments in algae

Algae are very diverse photosynthetic organisms, which differ from plants in that they are aquatic organisms, they do not present vascular tissue and do not generate an embryo. However, both types of organisms share the possession of photosynthetic pigments, which absorb and release energy that is later used by the cell. These pigments in addition to chlorophylls, are phycobiliproteins, fucoxanthins, xanthophylls and carotenes, which serve to trap the energy of light and lead it to the primary pigment, which is responsible for initiating oxygenic photosynthesis reactions. Algal phototrophs such as

dinoflagellates The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...

use

peridinin Peridinin is a light-harvesting apocarotenoid, a pigment associated with chlorophyll and found in the peridinin-chlorophyll-protein (PCP) light-harvesting complex in dinoflagellates, best studied in ''Amphidinium carterae''. Biological significan ...

as a light harvesting pigment. While carotenoids can be found complexed within chlorophyll-binding proteins such as the

photosynthetic reaction center A photosynthetic reaction center is a complex of several proteins, pigments and other co-factors that together execute the primary energy conversion reactions of photosynthesis. Molecular excitations, either originating directly from sunlight or t ...

s and

light-harvesting complexes A light-harvesting complex consists of a number of chromophores which are complex subunit proteins that may be part of a larger super complex of a photosystem, the functional unit in photosynthesis. It is used by plants and anoxygenic photosynthesi ...

, they also are found within dedicated carotenoid proteins such as the orange carotenoid protein of cyanobacteria.

Pigments in bacteria

Bacteria produce pigments such as carotenoids, melanin, violacein, prodigiosin, pyocyanin, actinorhodin, and zeaxanthin.

Cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...

produce phycocyanin, phycoerythrin, scytonemine, chlorophyll a, chlorophyll d, and chlorophyll f. Purple sulfur bacteria produce bacteriochlorophyll a and bacteriochlorophyll b. In cyanobacteria, many other carotenoids exist such as

myxoxanthophyll Myxoxanthophyll is a carotenoid glycoside pigment present (usually as rhamnosides) in the photosynthetic apparatus of cyanobacteria. It is named after the word "''Myxophyceae"'', a former term for cyanobacteria. As a monocyclic xanthophyll, it has ...

, synechoxanthin, and

echinenone Echinenone is a xanthophyll, with formula C40H54O. It is found in some cyanobacteria. It is synthesized from β-carotene by the enzyme beta-carotene ketolase (or CrtW). It has also been isolated from sea urchins Sea urchins () are spiny, g ...

Pigments in animals

Pigmentation is used by many animals for protection, by means of

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 ...

mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...

, or

warning coloration 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 ...

. Some animals including fish, amphibians and cephalopods use pigmented

s to provide camouflage that varies to match the background. Pigmentation is used in

signalling In signal processing, a signal is a function that conveys information about a phenomenon. Any quantity that can vary over space or time can be used as a signal to share messages between observers. The ''IEEE Transactions on Signal Processing'' ...

between animals, such as in courtship and reproductive behavior. For example, some

cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...

s use their chromatophores to communicate. The

photopigment Photopigments are unstable pigments that undergo a chemical change when they absorb light. The term is generally applied to the non-protein chromophore moiety of photosensitive chromoproteins, such as the pigments involved in photosynthesis and p ...

intercepts light as the first step in the perception of light. Skin pigments such as

may protect tissues from

sunburn Sunburn is a form of radiation burn that affects living tissue, such as skin, that results from an overexposure to ultraviolet (UV) radiation, usually from the Sun. Common symptoms in humans and animals include: red or reddish skin that is ho ...

ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...

radiation. However, some biological pigments in animals, such as

heme Heme, or haem (pronounced / hi:m/ ), is a precursor to hemoglobin, which is necessary to bind oxygen in the bloodstream. Heme is biosynthesized in both the bone marrow and the liver. In biochemical terms, heme is a coordination complex "consisti ...

groups that help to carry oxygen in the blood, are colored as a result of happenstance. Their color does not have a protective or signalling function.

Diseases and conditions

A variety of diseases and abnormal conditions that involve pigmentation are in humans and animals, either from absence of or loss of pigmentation or pigment cells, or from the excess production of pigment. *

Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and pink or blue eyes. Individuals with the condition are referred to as albino. Varied use and interpretation of the term ...

is an inherited disorder characterized by total or partial loss of

. Humans and animals that suffer from albinism are called "albinistic" (the term "albino" is also sometimes used, but may be considered offensive when applied to people). *

Lamellar ichthyosis Lamellar ichthyosis, also known as ichthyosis lamellaris and nonbullous congenital ichthyosis, is a rare inherited skin disorder, affecting around 1 in 600,000 people. Presentation Affected babies are born in a collodion membrane, a shiny, waxy- ...

, also called "fish scale disease", is an inherited condition in which one symptom is excess production of melanin. The skin is darker than normal, and is characterized by darkened, scaly, dry patches. *

Melasma Melasma (also known as chloasma faciei,James, William; Berger, Timothy; Elston, Dirk (2005). ''Andrews' Diseases of the Skin: Clinical Dermatology''. (10th ed.). Saunders. . or the mask of pregnancy when present in pregnant women) is a tan or dark ...

is a condition in which dark brown patches of pigment appear on the face, influenced by hormonal changes. When it occurs during a pregnancy, this condition is called ''the mask of pregnancy''. * ''ocular pigmentation'' is an accumulation of pigment in the eye, and may be caused by

latanoprost Latanoprost, sold under the brand name Xalatan among others, is a medication used to treat increased pressure inside the eye. This includes ocular hypertension and open angle glaucoma. It is applied as eye drops to the eyes. Onset of effects i ...

medication. Page 146 *

Vitiligo Vitiligo is a disorder that causes the skin to lose its color. Specific causes are unknown but studies suggest a link to immune system changes. Signs and symptoms The only sign of vitiligo is the presence of pale patchy areas of depigmen ...

is a condition in which there is a loss of pigment-producing cells called

melanocytes Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and heart. ...

in patches of skin.

Pigments in marine animals

Carotenoids and carotenoproteins

s are the most common group of pigments found in nature. Over 600 different kinds of carotenoids are found in animals, plants, and microorganisms. Animals are incapable of making their own carotenoids and thus rely on plants for these pigments. Carotenoproteins are especially common among marine animals. These complexes are responsible for the various colors (red, purple, blue, green, etc.) to these marine invertebrates for mating rituals and camouflage. There are two main types of carotenoproteins: Type A and Type B. Type A has carotenoids (chromogen) which are stoichiometrically associated with a simple protein (glycoprotein). The second type, Type B, has carotenoids which are associated with a lipo protein and is usually less stable. While Type A is commonly found in the surface (shells and skins) of marine invertebrates, Type B is usually in eggs, ovaries, and blood. The colors and characteristic absorption of these carotenoprotein complexes are based upon the chemical binding of the chromogen and the protein subunits. For example, the blue carotenoprotein, linckiacyanin has about 100-200 carotenoid molecules per every complex. In addition, the functions of these pigment-protein complexes also change their chemical structure as well. Carotenoproteins that are within the photosynthetic structure are more common, but complicated. Pigment-protein complexes that are outside of the photosynthetic system are less common, but have a simpler structure. For example, there are only two of these blue astaxanthin-proteins in the jellyfish, ''Velella velella'', contains only about 100 carotenoids per complex. A common carotenoid in animals is

astaxanthin Astaxanthin is a keto- carotenoid within a group of chemical compounds known as terpenes. Astaxanthin is a metabolite of zeaxanthin and canthaxanthin, containing both hydroxyl and ketone functional groups. It is a lipid-soluble pigment with r ...

, which gives off a purple-blue and green pigment. Astaxanthin's color is formed by creating complexes with proteins in a certain order. For example, the crustochrin has approximately 20 astaxanthin molecules bonded with protein. When the complexes interact by exciton-exciton interaction, it lowers the absorbance maximum, changing the different color pigments. In lobsters, there are various types of astaxanthin-protein complexes present. The first one is

crustacyanin Crustacyanin is a carotenoprotein biological pigment found in the exoskeleton of lobsters and responsible for their blue colour. β-Crustacyanin (β-CR), is composed of two stacked astaxanthin carotenoids that absorb at λ = 580–590 nm (2.1 ...

(max 632 nm), a slate-blue pigment found in the lobster's carapace. The second one is crustochrin (max 409), a yellow pigment which is found on the outer layer of the carapace. Lastly, the lipoglycoprotein and ovoverdin forms a bright green pigment that is usually present in the outer layers of the carapace and the lobster eggs

Tetrapyrroles

Tetrapyrroles Tetrapyrroles are a class of chemical compounds that contain four pyrrole or pyrrole-like rings. The pyrrole/pyrrole derivatives are linked by ( =- or -- units), in either a linear or a cyclic fashion. Pyrroles are a five-atom ring with four car ...

are the next most common group of pigments. They have four pyrrole rings, each ring consisting of C₄H₄NH. The main role of the tetrapyrroles is their connection in the biological oxidation process. Tetrapyrroles have a major role in electron transport and act as a replacement for many enzymes. They also have a role in the pigmentation of the marine organism's tissues.

Melanin

Melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...

is a class of compounds that serves as a pigment with different structures responsible for dark, tan, yellowish / reddish pigments in marine animals. It is produced as the amino acid tyrosine is converted into melanin, which is found in the skin, hair, and eyes. Derived from aerobic oxidation of phenols, they are polymers. There are several different types of melanins considering that they are an aggregate of smaller component molecules, such as nitrogen containing melanins. There are two classes of pigments: black and brown insoluble eumelanins, which are derived from aerobic oxidation of tyrosine in the presence of tyrosinase, and the alkali-soluble phaeomelanins which range from a yellow to red brown color, arising from the deviation of the eumelanin pathway through the intervention of cysteine and/or glutathione. Eumelanins are usually found in the skin and eyes. Several different melanins include melanoprotein (dark brown melanin that is stored in high concentrations in the ink sac of the cuttlefish Sepia Officianalis), echinoidea (found in sand dollars, and the hearts of sea urchins), holothuroidea (found in sea cucumbers), and ophiuroidea (found in brittle and snake stars). These melanins are possibly polymers which arise from the repeated coupling of simple bi-polyfunctional monomeric intermediates, or of high molecular weights. The compounds benzothiazole and tetrahydroisoquinoline ring systems act as UV-absorbing compounds.

Bioluminescence

The only light source in the deep sea, marine animals give off visible light energy called

bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some b ...

,Webexhibits.
Bioluminescence , Causes of Color
" WebExhibits. Web. 2 June 2010. a subset of

chemiluminescence Chemiluminescence (also chemoluminescence) is the emission of light (luminescence) as the result of a chemical reaction. There may also be limited emission of heat. Given reactants A and B, with an excited intermediate ◊, : + -> lozenge -> ...

. This is the chemical reaction in which chemical energy is converted to light energy. It is estimated that 90% of deep-sea animals produce some sort of bioluminescence. Considering that a large proportion of the visible light spectrum is absorbed before reaching the deep sea, most of the emitted light from the sea-animals is blue and green. However, some species may emit a red and infrared light, and there has even been a genus that is found to emit yellow bioluminescence. The organ that is responsible for the emission of bioluminescence is known as photophores. This type is only present in squid and fish, and is used to illuminate their ventral surfaces, which disguise their silhouettes from predators. The uses of the photophores in the sea-animals differ, such as lenses for controlling intensity of color, and the intensity of the light produced. Squids have both photophores and chromatophores which controls both of these intensities. Another thing that is responsible for the emission of bioluminescence, which is evident in the bursts of light that

jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...

emit, start with a luciferin (a photogen) and ends with the light emitter (a photagogikon.) Luciferin, luciferase, salt, and oxygen react and combine to create a single unit called photo-proteins, which can produce light when reacted with another molecule such as Ca+. Jellyfish use this as a defense mechanism; when a smaller predator is attempting to devour a jellyfish, it will flash its lights, which would therefore lure a larger predator and chase the smaller predator away. It is also used as mating behavior. In reef-building coral and sea anemones, they fluoresce; light is absorbed at one wavelength, and re-emitted at another. These pigments may act as natural sunscreens, aid in photosynthesis, serve as warning coloration, attract mates, warn rivals, or confuse predators.

Chromatophores

Chromatophore Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...

s are color pigment changing cells that are directly stimulated by central motor neurons. They are primarily used for quick environmental adaptation for camouflaging. The process of changing the color pigment of their skin relies on a single highly developed chromatophore cell and many muscles, nerves, glial and sheath cells. Chromatophores contract and contain vesicles that stores three different liquid pigments. Each color is indicated by the three types of chromatophore cells:

erythrophores Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contrast, ...

melanophores Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contrast, ...

, and

xanthophores Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contr ...

. The first type is the erythrophores, which contains reddish pigments such as carotenoids and pteridines. The second type is the melanophores, which contains black and brown pigments such as the melanins. The third type is the xanthophores which contains yellow pigments in the forms of carotenoids. The various colors are made by the combination of the different layers of the chromatophores. These cells are usually located beneath the skin or scale the animals. There are two categories of colors generated by the cell – biochromes and schematochromes. Biochromes are colors chemically formed microscopic, natural pigments. Their chemical composition is created to take in some color of light and reflect the rest. In contrast, schematochromes (structural colors) are colors created by light reflections from a colorless surface and refractions by tissues. Schematochromes act like prisms, refracting and dispersing visible light to the surroundings, which will eventually reflect a specific combination of colors. These categories are determined by the movement of pigments within the chromatophores. The physiological color changes are short-term and fast, found in fishes, and are a result from an animal's response to a change in the environment. In contrast, the morphological color changes are long-term changes, occurs in different stages of the animal, and are due to the change of numbers of chromatophores. To change the color pigments, transparency, or opacity, the cells alter in form and size, and stretch or contract their outer covering.

Photo-protective pigments

Due to damage from UV-A and UV-B, marine animals have evolved to have compounds that absorb UV light and act as sunscreen. Mycosporine-like amino acids (MAAs) can absorb UV rays at 310-360 nm. Melanin is another well-known UV-protector. Carotenoids and photopigments both indirectly act as photo-protective pigments, as they quench oxygen free-radicals. They also supplement photosynthetic pigments that absorb light energy in the blue region.

Defensive role of pigments

It's known that animals use their color patterns to warn off predators, however it has been observed that a sponge pigment mimicked a chemical which involved the regulation of moulting of an amphipod that was known to prey on sponges. So whenever that amphipod eats the sponge, the chemical pigments prevents the moulting, and the amphipod eventually dies.

Environmental influence on color

Coloration in invertebrates varies based on the depth, water temperature, food source, currents, geographic location, light exposure, and sedimentation. For example, the amount of carotenoid a certain sea anemone decreases as we go deeper into the ocean. Thus, the marine life that resides on deeper waters is less brilliant than the organisms that live in well-lit areas due to the reduction of pigments. In the colonies of the colonial ascidian-cyanophyte symbiosis Trididemnum solidum, their colors are different depending on the light regime in which they live. The colonies that are exposed to full sunlight are heavily calcified, thicker, and are white. In contrast the colonies that live in shaded areas have more phycoerythrin (pigment that absorbs green) in comparison to phycocyanin (pigment that absorbs red), thinner, and are purple. The purple color in the shaded colonies are mainly due to the phycobilin pigment of the algae, meaning the variation of exposure in light changes the colors of these colonies.

Adaptive coloration

Aposematism is the warning coloration to signal potential predators to stay away. In many chromodrorid nudibranchs, they take in distasteful and toxic chemicals emitted from sponges and store them in their repugnatorial glands (located around the mantle edge). Predators of nudibranchs have learned to avoid these certain nudibranchs based on their bright color patterns. Preys also protect themselves by their toxic compounds ranging from a variety of organic and inorganic compounds.

Physiological activities

Pigments of marine animals serve several different purposes, other than defensive roles. Some pigments are known to protect against UV (see photo-protective pigments.) In the nudibranch Nembrotha Kubaryana, tetrapyrrole pigment 13 has been found to be a potent antimicrobial agent. Also in this creature, tamjamines A, B, C, E, and F has shown antimicrobial, antitumor, and immunosuppressive activities. Sesquiterpenoids are recognized for their blue and purple colors, but it has also been reported to exhibit various bioactivities such as antibacterial, immunoregulating, antimicrobial, and cytotoxic, as well as the inhibitory activity against cell division in the fertilized sea urchin and ascidian eggs. Several other pigments have been shown to be cytotoxic. In fact, two new carotenoids that were isolated from a sponge called Phakellia stelliderma showed mild cytotoxicity against mouse leukemia cells. Other pigments with medical involvements include

scytonemin Scytonemin is a secondary metabolite and an Extracellular matrix, extracellular matrix (sheath) Biological pigment, pigment synthesized by many strains of cyanobacteria, including ''Nostoc'', ''Scytonema'', Calothrix crustacea, ''Calothrix'', ''Lyn ...

, topsentins, and debromohymenialdisine have several lead compounds in the field of inflammation, rheumatoid arthritis and osteoarthritis respectively. There's evidence that topsentins are potent mediators of immunogenic inflation, and topsentin and scytonemin are potent inhibitors of neurogenic inflammation.

Uses

Pigments may be extracted and used as

dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...

s. Pigments (such as astaxanthin and lycopene) are used as dietary supplements.

References

External links

* * {{Authority control Biological pigments Warning coloration

Biological pigments

Pigments in plants

Pigments in algae

Pigments in bacteria

Pigments in animals

Diseases and conditions

Pigments in marine animals

Carotenoids and carotenoproteins

Tetrapyrroles

Melanin

Bioluminescence

Chromatophores

Photo-protective pigments

Defensive role of pigments

Environmental influence on color

Adaptive coloration

Physiological activities

Uses

See also

References

External links