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Biological pigment
Biological pigments, also known simply as pigments or
biochromes,[1] are substances produced by living organisms that
have a color resulting from selective color absorption. Biological
pigments include plant pigments and flower pigments. Many
biological structures, such as skin, eyes, feathers, fur and hair
contain pigments such as melanin in specialized cells called
chromatophores. In some species, pigments accrue over very long
periods during an individual's lifespan.[2]

Pigment color differs from structural color in that it is the same for
all viewing angles, whereas structural color is the result of selective
reflection or iridescence, usually because of multilayer structures.
For example, butterfly wings typically contain structural color,
although many butterflies have cells that contain pigment as well.[3]
The budgerigar gets its yellow
color from a psittacofulvin
pigment and its green color from
Contents a combination of the same
yellow pigment and blue
Biological pigments
structural color. The blue and
Pigments in plants white bird in the background
Pigments in algae lacks the yellow pigment. The
dark markings on both birds are
Pigments in bacteria due to the black pigment
Pigments in animals eumelanin.
Diseases and conditions
Pigments in marine animals
Uses
See also
References
External links

Biological pigments
See conjugated systems for electron bond chemistry that causes these molecules to have pigment.

Heme/porphyrin-based: chlorophyll, bilirubin, hemocyanin, hemoglobin, myoglobin

Light-emitting: luciferin
Carotenoids:
Hematochromes (algal pigments, mixes of carotenoids and their derivates)
Carotenes: alpha and beta carotene, lycopene, rhodopsin
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Xanthophylls: canthaxanthin, zeaxanthin, lutein

Proteinaceous: phytochrome, phycobiliproteins
Psittacofulvins: a class of red and yellow pigments unique to parrots
Turacin and Turacoverdin: red and green pigments found in turacos and related species
Other: melanin, urochrome, flavonoids

Pigments in plants
The primary function of pigments in plants is photosynthesis,
which uses the green pigment chlorophyll and several colorful
pigments that absorb as much light energy as possible.[4][5]
Pigments are also known to play a role in pollination where
pigment accumulation or loss can lead to floral color change,
signaling to pollinators which flowers are rewarding and contain
more pollen and nectar.[6]
Space-filling model of the
Plant pigments include many molecules, such as porphyrins,
chlorophyll molecule.
carotenoids, anthocyanins and betalains. All biological pigments
selectively absorb certain wavelengths of light while reflecting
others.[4][5]

The principal pigments responsible are:

Chlorophyll is the primary pigment in plants; it is a chlorin

that absorbs blue and red wavelengths of light while reflecting
a majority of green. It is the presence and relative abundance
of chlorophyll that gives plants their green color. All land
plants and green algae possess two forms of this pigment:
chlorophyll a and chlorophyll b. Kelps, diatoms, and other Anthocyanin gives these pansies
photosynthetic heterokonts contain chlorophyll c instead of b, their purple pigmentation.
while red algae possess only chlorophyll a. All chlorophylls
serve as the primary means plants use to intercept light in
order to fuel photosynthesis.
Carotenoids are red, orange, or yellow tetraterpenoids. During the process of photosynthesis,
they have functions in light-harvesting (as accessory pigments), in photoprotection (energy
dissipation via non-photochemical quenching 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.
Betalains are red or yellow pigments. Like anthocyanins they are water-soluble, but unlike
anthocyanins they are synthesized from tyrosine. This class of pigments is found only in the
Caryophyllales (including cactus and amaranth), and never co-occur in plants with
anthocyanins.[5] Betalains are responsible for the deep red color of beets.
Anthocyanins (literally "flower blue") are water-soluble flavonoid pigments 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 petals of flowers of many species.[5]

Plants, in general, contain six ubiquitous carotenoids: neoxanthin, violaxanthin, antheraxanthin,

zeaxanthin, lutein and β-carotene.[7] Lutein is a yellow pigment found in fruits and vegetables and is
the most abundant carotenoid in plants. Lycopene is the red pigment responsible for the color of
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tomatoes. Other less common carotenoids in plants include lutein epoxide (in many woody species),
lactucaxanthin (found in lettuce), and alpha carotene (found in carrots).[8]

A particularly noticeable manifestation of pigmentation in plants

is seen with autumn leaf color, a phenomenon that affects the
normally green leaves of many deciduous trees and shrubs
whereby they take on, during a few weeks in the autumn season,
various shades of red, yellow, purple, and brown.[9]

Chlorophylls degrade into colorless tetrapyrroles known as

nonfluorescent chlorophyll catabolites (NCCs).[10]
As the
predominant chlorophylls degrade, the hidden pigments of yellow
xanthophylls and orange beta-carotene are revealed. These
pigments are present throughout the year, but the red pigments,
the anthocyanins, are synthesized de novo once roughly half of
chlorophyll has been degraded. The amino acids released from
degradation of light harvesting complexes are stored all winter in
the tree's roots, branches, stems, and trunk until next spring
when they are recycled to re‑leaf the tree.
Bougainvillea bracts get their color
Pigments in algae from betalains

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 use peridinin as a light harvesting pigment. While
carotenoids can be found complexed within chlorophyll-binding proteins such as the photosynthetic
reaction centers and light-harvesting complexes, they also are found within dedicated carotenoid
proteins such as the orange carotenoid protein of cyanobacteria.

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Table 1. Pigments in algae

Group Pigment
Green algae -Chlorophyll a and b
-Chlorophyll a

-Ficobiliproteins
Red algae
-Phycoerythrin red-purple pigment, it is the dominant one in the species that
have a red-purple color.

-Chlorophyll a and c

Golden and Brown -Xantophyll

algae
-Fucoxantin

Pigments in bacteria
Bacteria produce pigments such as carotenoids,
melanin, violacein, prodigiosin, pyocyanin,
actinorhodin, and zeaxanthin. Cyanobacteria produce
phycocyanin, phycoerythrin, scytonemine, chlorophyll
a, chlorophyll d, and chlorophyll f. Purple sulfur
bacteria produce bacteriochlorophyll a and
bacteriochlorophyll b.[11] In cyanobacteria, many other
carotenoids exist such as canthaxanthin,
myxoxanthophyll, synechoxanthin, and echinenone.

Cyanobacteria exhibit different pigments

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Table 2. Pigments in bacteria

Group Pigment
-Chlorophyll a, Chlorophyll d and Chlorophyll f

-Phycocyanin blue-green pigment, is the dominant pigment in cyanobacteria

-Allophycocyanin

-Phycoerithrin
Cyanobacteria
-Scytonemin

-Cyanophycin

-Carotenoids

Purple bacteria -Bacteriochlorophyll a and b

Green bacteria -Bacteriochlorophyll c and e
Chromobacterium -Violacein
Streptomyces -Melanin
Micromonospora -Anthraquinone

Pigments in animals
Most of the hemoglobin is removed when animals are slaughtered and bled. Thus, in properly bled
muscle tissue myoglobin is responsible for 90% or more of the pigmentation. The myoglobin quantity
varies considerably among muscle tissues and is influenced by species, age, sex, and physical activity.

Pigmentation is used by many animals for protection, by means of camouflage, mimicry, or warning
coloration. Some animals including fish, amphibians and cephalopods use pigmented
chromatophores to provide camouflage that varies to match the background.

Pigmentation is used in signalling between animals, such as in courtship and reproductive behavior.
For example, some cephalopods use their chromatophores to communicate.

The photopigment rhodopsin intercepts light as the first step in the perception of light.

Skin pigments such as melanin may protect tissues from sunburn by ultraviolet radiation.

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

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Albinism is an inherited disorder characterized by total or partial loss of melanin. 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, 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 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
medication.[12]
Vitiligo is a condition in which there is a loss of pigment-producing cells called melanocytes in
patches of skin.

Pigments in marine animals

Carotenoids and carotenoproteins

Carotenoids are the most common group of pigments found in nature.[13] 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.[14] 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, 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 (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[15][16]
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Tetrapyrroles

Tetrapyrroles 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[17] 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,[18] a subset of chemiluminescence. 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 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.

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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

Chromatophores 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, melanophores, and xanthophores. 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
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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, 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 dyes.

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

See also
Photosynthetic pigment

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External links
Ernest Ingersoll (1920). "Color in Plants"  (https://en.wikisource.org/wiki/The_Encyclopedia_Ameri
cana_(1920)/Color_in_Plants). Encyclopedia Americana.
John Merle Coulter (1905). "Color, in Plants"  (https://en.wikisource.org/wiki/The_New_Internation
al_Encyclop%C3%A6dia/Color,_in_Plants). New International Encyclopedia.

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