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Trichromacy or trichromatism is the possession of three independent channels for conveying
color Color (or colour in English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is the visual perception based on the electromagnetic spectrum. Though co ...
information, derived from the three different types of
cone cell Cone cells or cones are photoreceptor cells in the retina of the vertebrate eye. Cones are active in daylight conditions and enable photopic vision, as opposed to rod cells, which are active in dim light and enable scotopic vision. Most v ...
s in the
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
. Organisms with trichromacy are called trichromats. The normal explanation of trichromacy is that the organism's
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
contains three types of color receptors (called
cone cell Cone cells or cones are photoreceptor cells in the retina of the vertebrate eye. Cones are active in daylight conditions and enable photopic vision, as opposed to rod cells, which are active in dim light and enable scotopic vision. Most v ...
s in
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s) with different absorption spectra. In actuality, the number of such receptor types may be greater than three, since different types may be active at different light intensities. In vertebrates with three types of cone cells, at low light intensities the
rod cell Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in ...
s may contribute to
color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different frequencies independently of light intensity. Color perception is a part of the larger visual system and is mediated by a co ...
.


Humans and other animals that are trichromats

Human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s and some other
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s have evolved trichromacy based partly on
pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
s inherited from early vertebrates. In fish and birds, for example, four pigments are used for vision. These extra cone receptor visual pigments detect energy of other
wavelength In physics and mathematics, wavelength or spatial period of a wave or periodic function is the distance over which the wave's shape repeats. In other words, it is the distance between consecutive corresponding points of the same ''phase (waves ...
s, sometimes including
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
. Eventually two of these pigments were lost (in placental mammals) and another was gained, resulting in trichromacy among some
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s. Humans and closely related primates are usually trichromats, as are some of the females of most species of
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboi ...
s, and both male and female
howler monkey Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropical realm, Neotropics and are among the largest of the New World monkey, platyrrhines along with the muriquis (''Brachyte ...
s. Recent research suggests that trichromacy may also be quite general among
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s. A study conducted regarding trichromacy in
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
n marsupials suggests the medium wavelength sensitivity (MWS), cones of the honey possum (''Tarsipes rostratus'') and the fat-tailed dunnart (''Sminthopsis crassicaudata'') are features coming from the inherited reptilian
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
l arrangement. Another study used behavioral tests, genetic analyses, and immunohistochemistry and found trichromacy and ultraviolet vision in nocturnal sugar gliders (''Petaurus breviceps''). The possibility of trichromacy in marsupials potentially has another
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
ary basis than that of
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s. Further
biological Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, origin, evolution, and distribution of ...
and
behavioural Behavior (American English) or behaviour (British English) is the range of actions of individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as well as the inanimate ph ...
tests may verify if trichromacy is a common characteristic of marsupials. Most other mammals are currently thought to be
dichromat Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats require ...
s, with only two types of cone (though limited trichromacy is possible at low light levels where the rods and cones are both active). Most studies of carnivores, as of other mammals, reveal
dichromacy Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats requir ...
; examples include the domestic
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers. ...
, the
ferret The ferret (''Mustela furo'') is a small, domesticated species belonging to the family Mustelidae. The ferret is most likely a domesticated form of the wild European polecat (''Mustela putorius''), as evidenced by the ferret's ability to inter ...
, and the
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
. Some species of
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s (such as
honeybee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the c ...
s) are also trichromats, being sensitive to
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
, blue and green instead of blue, green and red. Research indicates that trichromacy allows animals to distinguish brightly colored fruit and young leaves from other vegetation that is not beneficial to their survival. Another theory is that detecting skin flushing and thereby mood may have influenced the development of primate trichromate vision. The color red also has other effects on primate and human behavior as discussed in the color psychology article.Diana Widermann, Robert A. Barton, and Russel A. Hill. Evolutionary perspectives on sport and competition. In


Types of cones specifically found in primates

Primates are the only known placental mammalian trichromats. Their eyes include three different kinds of cones, each containing a different
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 (chemistry), moiety of photosensitive chromoproteins, such as the pigments involved in ph ...
(
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become retinylidene proteins, but are usually still called opsins regardless. Most pro ...
). Their peak sensitivities lie in the blue (short-wavelength S cones), green (medium-wavelength M cones) and yellow-green (long-wavelength L cones) regions of the color spectrum. S cones make up 5–10% of the cones and form a regular mosaic. Special bipolar and
ganglion A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there are ...
cells pass those signals from S cones and there is evidence that they have a separate signal pathway through the
thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...
to the
visual cortex The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalam ...
as well. On the other hand, the L and M cones are hard to distinguish by their shapes or other anatomical means – their opsins differ in only 15 out of 363 amino acids, so no one has yet succeeded in producing specific antibodies to them. But Mollon and Bowmaker did find that L cones and M cones are randomly distributed and are in equal numbers.


Mechanism of trichromatic color vision

Trichromatic color vision is the ability of humans and some other animals to see different
color Color (or colour in English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is the visual perception based on the electromagnetic spectrum. Though co ...
s, mediated by interactions among three types of color-sensing
cone cell Cone cells or cones are photoreceptor cells in the retina of the vertebrate eye. Cones are active in daylight conditions and enable photopic vision, as opposed to rod cells, which are active in dim light and enable scotopic vision. Most v ...
s. The '' trichromatic color theory'' began in the 18th century, when Thomas Young proposed that color vision was a result of three different
photoreceptor cell A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiation ...
s. From the middle of the 19th century, in his ''Treatise on Physiological Optics'',
Hermann von Helmholtz Hermann Ludwig Ferdinand von Helmholtz (; ; 31 August 1821 – 8 September 1894; "von" since 1883) was a German physicist and physician who made significant contributions in several scientific fields, particularly hydrodynamic stability. The ...
later expanded on Young's ideas using color-matching experiments which showed that people with normal vision needed three wavelengths to create the normal range of colors. Physiological evidence for trichromatic theory was later given by Gunnar Svaetichin (1956). Each of the three types of cones in the
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
of the
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
contains a different type of photosensitive pigment, which is composed of a
transmembrane protein A transmembrane protein is a type of integral membrane protein that spans the entirety of the cell membrane. Many transmembrane proteins function as gateways to permit the transport of specific substances across the membrane. They frequently un ...
called
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become retinylidene proteins, but are usually still called opsins regardless. Most pro ...
and a light-sensitive molecule called 11-cis retinal. Each different pigment is especially sensitive to a certain
wavelength In physics and mathematics, wavelength or spatial period of a wave or periodic function is the distance over which the wave's shape repeats. In other words, it is the distance between consecutive corresponding points of the same ''phase (waves ...
of
light Light, visible light, or visible radiation is electromagnetic radiation that can be visual perception, perceived by the human eye. Visible light spans the visible spectrum and is usually defined as having wavelengths in the range of 400– ...
(that is, the pigment is most likely to produce a cellular response when it is hit by a
photon A photon () is an elementary particle that is a quantum of the electromagnetic field, including electromagnetic radiation such as light and radio waves, and the force carrier for the electromagnetic force. Photons are massless particles that can ...
with the specific wavelength to which that pigment is most sensitive). The three types of cones are L, M, and S, which have pigments that respond best to light of long (especially 560 nm), medium (530 nm), and short (420 nm) wavelengths respectively. Since the likelihood of response of a given cone varies not only with the ''wavelength'' of the light that hits it but also with its ''intensity'', the
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
would not be able to discriminate different colors if it had input from only one type of cone. Thus, interaction between at least two types of cone is necessary to produce the ability to perceive color. With at least two types of cones, the brain can compare the signals from each type and determine both the intensity and color of the light. For example, moderate stimulation of a medium-wavelength cone cell could mean that it is being stimulated by very bright red (long-wavelength) light, or by not very intense yellowish-green light. But very bright red light would produce a stronger response from L cones than from M cones, while not very intense yellowish light would produce a stronger response from M cones than from other cones. Thus trichromatic color vision is accomplished by using combinations of cell responses. It is estimated that the average human can distinguish up to ten million different colors.


See also

*
Visual system The visual system is the physiological basis of visual perception (the ability to perception, detect and process light). The system detects, phototransduction, transduces and interprets information concerning light within the visible range to ...
*
Monochromacy Monochromacy (from Greek ''mono'', meaning "one" and ''chromo'', meaning "color") is the ability of organisms to perceive only light intensity without respect to spectral composition. Organisms with monochromacy lack color vision and can only ...
*
Dichromacy Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats requir ...
*
Tetrachromacy Tetrachromacy (from Ancient Greek ''tetra'', meaning "four" and ''chroma'', meaning "color") is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms ...
* Pentachromacy *
Mantis shrimp Mantis shrimp are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 400 million years ago, with more than 520 extant species of mantis shrimp known. All li ...
(dodecachromats) * Evolution of color vision in primates *
Young–Helmholtz theory The Young–Helmholtz theory (based on the work of Thomas Young and Hermann von Helmholtz in the 19th century), also known as the trichromatic theory, is a theory of trichromatic color vision – the manner in which the visual system gives rise ...
*
LMS color space LMS (long, medium, short), is a color space which represents the response of the three types of Cone cell, cones of the human eye, named for their responsivity (sensitivity) peaks at long, medium, and short wavelengths. The numerical range is ...


References


External links


The Straight Dope
"Are cats and dogs really color-blind? How do they know?" * {{color vision Color vision Visual system