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Trichromacy or trichromatism is the possessing of three independent channels for conveying
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 assoc ...
information, derived from the three different types of
cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Con ...
s in the
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 ...
. Organisms with trichromacy are called trichromats. The normal explanation of trichromacy is that the organism's
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
contains three types of color receptors (called
cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Con ...
s in
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, wi ...
) 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 p ...
s may contribute to
color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of ...
.


Humans and other animals that are trichromats

Human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s and some other
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...
have evolved trichromacy based partly on
pigment 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 comp ...
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, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tr ...
s, sometimes including
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiati ...
. Eventually two of these pigments were lost (in placental mammals) and another was gained, resulting in trichromacy among some
primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...
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 Ceboid ...
s, and both male and female
howler monkey Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropics and are among the largest of the platyrrhines along with the muriquis (''Brachyteles''), the spider monkeys (''Atel ...
s. Recent research suggests that trichromacy may also be quite general among
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a ...
s. A study conducted regarding trichromacy in Australian marsupials suggests the medium wavelength sensitivity (MWS), cones of the
honey possum The honey possum or noolbenger (''Tarsipes rostratus''), is a tiny species of marsupial that feeds on the nectar and pollen of a diverse range of flowering plants. Found in southwest Australia, it is an important pollinator for such plants as '' ...
(''Tarsipes rostratus'') and the fat-tailed dunnart (''Sminthopsis crassicaudata'') are features coming from the inherited reptilian
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
l arrangement. The possibility of trichromacy in marsupials potentially has another
evolutionary Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
basis than that of
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter includin ...
. Further
biological Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
and behavioural 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 require ...
, examples including the domestic dog, 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''), evidenced by their interfertility. Other mu ...
, 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 ...
. Some species of
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
(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 Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cos ...
s) are also trichromats, being sensitive to
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiati ...
, 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 Color psychology is the study of hues as a determinant of human behavior. Color influences perceptions that are not obvious, such as the taste of food. Colors have qualities that can cause certain emotions in people. How color influences individ ...
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 (
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 Bipolar may refer to: Astronomy * Bipolar nebula, a distinctive nebular formation * Bipolar outflow, two continuous flows of gas from the poles of a star Mathematics * Bipolar coordinates, a two-dimensional orthogonal coordinate system * Bipolar ...
and ganglion cells pass those signals from S cones and there is evidence that they have a separate signal pathway through the
thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all direction ...
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 thalamus and ...
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 nobody 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 (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 assoc ...
s, mediated by interactions among three types of color-sensing
cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Con ...
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 radiati ...
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) was a German physicist and physician who made significant contributions in several scientific fields, particularly hydrodynamic stability. The Helmholtz Associatio ...
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 Gunnar Nils Toivo Svaetichin (13 January 1915 – 23 March 1981) was a Swedish-Finnish-Venezuelan physiologist who, in 1956, showed by examining the external layers of fish retinas that electroretinograms display particular sensitivity to three di ...
(1956). Each of the three types of cones in the
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
of the
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 ...
contains a different type of photosensitive pigment, which is composed of a
transmembrane protein A transmembrane protein (TP) 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 frequentl ...
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, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tr ...
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 te ...
(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 particle, massless ...
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 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 special ...
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 comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight ...
*
Monochromacy Monochromacy (from Greek ''mono'', meaning "one" and ''chromo'', meaning "color") is the ability of organisms or machines to perceive only light intensity, without respect to spectral composition (color). Organisms with monochromacy are called ...
*
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 require ...
*
Tetrachromacy Tetrachromacy (from Greek ''tetra'', meaning "four" and ''chromo'', 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 with tetr ...
* Pentachromacy *
Mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...
(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 cones of the human eye, named for their responsivity (sensitivity) peaks at long, medium, and short wavelengths. The numerical range is generall ...


References


External links


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