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Color vision, a feature of
visual perception Visual perception is the ability to interpret the surrounding environment through photopic vision (daytime vision), color vision, scotopic vision (night vision), and mesopic vision (twilight vision), using light in the visible spectrum ref ...
, 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 the larger visual system and is mediated by a complex process between neurons that begins with differential stimulation of different types of photoreceptors by light entering the eye. Those photoreceptors then emit outputs that are propagated through many layers of neurons and then ultimately to the
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
. Color vision is found in many animals and is mediated by similar underlying mechanisms with common types of biological molecules and a complex history of
evolution 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 ...
in different animal taxa. In
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 including ...
, color vision may have evolved under selective pressure for a variety of visual tasks including the foraging for nutritious young leaves, ripe fruit, and flowers, as well as detecting predator camouflage and emotional states in other primates.


Wavelength

Isaac Newton Sir Isaac Newton (25 December 1642 – 20 March 1726/27) was an English mathematician, physicist, astronomer, alchemist, Theology, theologian, and author (described in his time as a "natural philosophy, natural philosopher"), widely ...
discovered that
white light White is the lightest color and is achromatic (having no hue). It is the color of objects such as snow, chalk, and milk, and is the opposite of black. White objects fully reflect and scatter all the visible wavelengths of light. White on ...
after being split into its component colors when passed through a dispersive prism could be recombined to make white light by passing them through a different prism. The
visible light spectrum The visible spectrum is the portion of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called ''visible light'' or simply light. A typical human eye will respond to wavele ...
ranges from about 380 to 740 nanometers. Spectral colors (colors that are produced by a narrow band of wavelengths) such as red, orange, yellow, green, cyan, blue, and violet can be found in this range. These spectral colors do not refer to a single wavelength, but rather to a set of wavelengths: red, 625–740 nm; orange, 590–625 nm; yellow, 565–590 nm; green, 500–565 nm; cyan, 485–500 nm; blue, 450–485 nm; violet, 380–450 nm. Wavelengths longer or shorter than this range are called
infrared Infrared (IR), sometimes called infrared light, is electromagnetic radiation (EMR) with wavelengths longer than those of Light, visible light. It is therefore invisible to the human eye. IR is generally understood to encompass wavelengths from ...
or
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 radiation ...
, respectively. Humans cannot generally see these wavelengths, but other animals may.


Hue detection

Sufficient differences in wavelength cause a difference in the perceived hue; the just-noticeable difference in wavelength varies from about 1  nm in the blue-green and
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 th ...
wavelengths to 10 nm and more in the longer red and shorter blue wavelengths. Although the human eye can distinguish up to a few hundred hues, when those pure spectral colors are mixed together or diluted with white light, the number of distinguishable chromaticities can be quite high. In very low light levels, vision is scotopic: light is detected by rod cells of 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 the ...
. Rods are maximally sensitive to wavelengths near 500 nm and play little, if any, role in color vision. In brighter light, such as daylight, vision is photopic: light is detected by
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. Cone ...
s which are responsible for color vision. Cones are sensitive to a range of wavelengths, but are most sensitive to wavelengths near 555 nm. Between these regions, mesopic vision comes into play and both rods and cones provide signals to the
retinal ganglion cell A retinal ganglion cell (RGC) is a type of neuron located near the inner surface (the ganglion cell layer) of the retina of the eye. It receives visual information from photoreceptors via two intermediate neuron types: bipolar cells and reti ...
s. The shift in color perception from dim light to daylight gives rise to differences known as the Purkinje effect. The perception of "white" is formed by the entire spectrum of visible light, or by mixing colors of just a few wavelengths in animals with few types of color receptors. In humans, white light can be perceived by combining wavelengths such as red, green, and blue, or just a pair of
complementary color Complementary colors are pairs of colors which, when combined or mixed, cancel each other out (lose hue) by producing a grayscale color like white or black. When placed next to each other, they create the strongest contrast for those t ...
s such as blue and yellow.


Non-spectral colors

There are a variety of colors in addition to spectral colors and their hues. These include grayscale colors, shades of colors obtained by mixing grayscale colors with spectral colors, violet-red colors,
impossible color Impossible colors are colors that do not appear in ordinary visual functioning. Different color theories suggest different hypothetical colors that humans are incapable of perceiving for one reason or another, and fictional colors are rou ...
s, and
metallic color A metallic color is a color that appears to be that of a polished metal. The visual sensation usually associated with metals is its metallic shine. This cannot be reproduced by a simple solid color, because the shiny effect is due to the materi ...
s. Grayscale colors include white, gray, and black. Rods contain rhodopsin, which reacts to light intensity, providing grayscale coloring. Shades include colors such as pink or brown. Pink is obtained from mixing red and white. Brown may be obtain from mixing orange with gray or black. Navy is obtained from mixing blue and black. Violet-red colors include hues and shades of magenta. The light spectrum is a line on which violet is one end and the other is red, and yet we see hues of purple that connect those two colors. Impossible colors are a combination of cone responses that cannot be naturally produced. For example, medium cones cannot be activated completely on their own; if they were, we would see a 'hyper-green' color.


Dimensionality

Color vision is categorized foremost according to the dimensionality of the color
gamut In color reproduction, including computer graphics and photography, the gamut, or color gamut , is a certain ''complete subset'' of colors. The most common usage refers to the subset of colors which can be accurately represented in a given circ ...
, which is defined by the number of
primaries Primary elections, or direct primary are a voting process by which voters can indicate their preference for their party's candidate, or a candidate in general, in an upcoming general election, local election, or by-election. Depending on the c ...
required to represent the color vision. This is generally equal to the number of
photopsin Vertebrate visual opsins are a subclass of ciliary opsins and mediate vision in vertebrates. They include the opsins in human rod and cone cells. They are often abbreviated to ''opsin'', as they were the first opsins discovered and are still th ...
s expressed: a correlation that holds for
vertebrate 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, with ...
s but not
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s. The common vertebrate ancestor possessed four photopsins (expressed in cones) plus rhodopsin (expressed in rods), so was tetrachromatic. However, many vertebrate lineages have lost one or many photopsin genes, leading to lower-dimension color vision. The dimensions of color vision range from 1-dimensional and up: * Monochromacy - 1D color vision - lack of any color perception *
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 ...
- 2D color vision - dimensionality of most mammals and a quarter of
color blind Color blindness or color vision deficiency (CVD) is the decreased ability to see color or differences in color. It can impair tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights. Color blindness may make some aca ...
humans *
Trichromacy Trichromacy or trichromatism is the possessing of three independent channels for conveying color information, derived from the three different types of cone cells in the eye. Organisms with trichromacy are called trichromats. The normal expl ...
- 3D color vision - dimensionality of most humans *
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 te ...
- 4D color vision - dimensionality of most
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s and
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
* Pentachromacy and higher - 5D+ color vision - rare in vertebrates


Physiology of color perception

Perception of color begins with specialized retinal cells known as
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. Cone ...
s. Cone cells contain different forms of opsin – a pigment protein – that have different spectral sensitivities. Humans contain three types, resulting in trichromatic color vision. Each individual cone contains pigments composed of
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 ...
apoprotein covalently linked to a light-absorbing
prosthetic group A prosthetic group is the non-amino acid component that is part of the structure of the heteroproteins or conjugated proteins, being tightly linked to the apoprotein. Not to be confused with the cofactor that binds to the enzyme apoenzyme (eith ...
: either 11-''cis''-hydroretinal or, more rarely, 11-''cis''-dehydroretinal. The cones are conventionally labeled according to the ordering of the wavelengths of the peaks of their spectral sensitivities: short (S), medium (M), and long (L) cone types. These three types do not correspond well to particular colors as we know them. Rather, the perception of color is achieved by a complex process that starts with the differential output of these cells in the retina and which is finalized in 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 associative areas of the brain. For example, while the L cones have been referred to simply as red receptors, microspectrophotometry has shown that their peak sensitivity is in the greenish-yellow region of the spectrum. Similarly, the S cones and M cones do not directly correspond to
blue Blue is one of the three primary colours in the RYB colour model (traditional colour theory), as well as in the RGB (additive) colour model. It lies between violet and cyan on the spectrum of visible light. The eye perceives blue when ...
and
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 nm. In subtractive color systems, used in painting and color printing, it is created by a combin ...
, although they are often described as such. The
RGB color model The RGB color model is an additive color model in which the red, green and blue primary colors of light are added together in various ways to reproduce a broad array of colors. The name of the model comes from the initials of the three ad ...
, therefore, is a convenient means for representing color but is not directly based on the types of cones in the human eye. The peak response of human cone cells varies, even among individuals with so-called normal color vision; in some non-human species this polymorphic variation is even greater, and it may well be adaptive.


Theories

Two complementary theories of color vision are the trichromatic theory and the
opponent process The opponent process is a color theory that states that the human visual system interprets information about color by processing signals from photoreceptor cells in an antagonistic manner. The opponent-process theory suggests that there are thr ...
theory. The trichromatic theory, or Young–Helmholtz theory, proposed in the 19th century by Thomas Young and
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 Associat ...
, posits three types of cones preferentially sensitive to blue, green, and red, respectively. Ewald Hering proposed the opponent process theory in 1872. It states that the visual system interprets color in an antagonistic way: red vs. green, blue vs. yellow, black vs. white. Both theories are generally accepted as valid, describing different stages in visual physiology, visualized in the adjacent diagram. Green–magenta and blue—yellow are scales with mutually exclusive boundaries. In the same way that there cannot exist a "slightly negative" positive number, a single eye cannot perceive a bluish-yellow or a reddish-green. Although these two theories are both currently widely accepted theories, past and more recent work has led to criticism of the opponent process theory, stemming from a number of what are presented as discrepancies in the standard opponent process theory. For example, the phenomenon of an after-image of complementary color can be induced by fatiguing the cells responsible for color perception, by staring at a vibrant color for a length of time, and then looking at a white surface. This phenomenon of complementary colors demonstrates cyan, rather than green, to be the complement of red and magenta, rather than red, to be the complement of green, as well as demonstrating, as a consequence, that the reddish-green color proposed to be impossible by opponent process theory is, in fact, the color yellow. Although this phenomenon is more readily explained by the trichromatic theory, explanations for the discrepancy may include alterations to the opponent process theory, such as redefining the opponent colors as red vs. cyan, to reflect this effect. Despite such criticisms, both theories remain in use. A recent demonstration, using the Color Mondrian, has shown that, just as the color of a surface that is part of a complex 'natural' scene is independent of the wavelength-energy composition of the light reflected from it alone but depends upon the composition of the light reflected from its surrounds as well, so the after image produced by looking at a given part of a complex scene is also independent of the wavelength energy-composition of the light reflected from it alone. Thus, while the color of the after-image produced by looking at a green surface that is reflecting more "green" (middle-wave) than "red" (long-wave) light is magenta, so is the after image of the same surface when it reflects more "red" than "green" light (when it is still perceived as green). This would seem to rule out an explanation of color opponency based on retinal cone adaptation.


Cone cells in the human eye

A range of wavelengths of light stimulates each of these receptor types to varying degrees. The brain combines the information from each type of receptor to give rise to different perceptions of different wavelengths of light. Cones and rods are not evenly distributed in the human eye. Cones have a high density at the
fovea Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure. Human anatomy *Fovea centralis of the retina * Fovea buccalis or Dimple * Fovea of the femoral head * Trochlear fovea of the fr ...
and a low density in the rest of the retina. Thus color information is mostly taken in at the fovea. Humans have poor color perception in their peripheral vision, and much of the color we see in our periphery may be filled in by what our brains expect to be there on the basis of context and memories. However, our accuracy of color perception in the periphery increases with the size of stimulus. The opsins (photopigments) present in the L and M cones are encoded on the X
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
; defective encoding of these leads to the two most common forms of color blindness. The '' OPN1LW'' gene, which encodes the opsin present in the L cones, is highly polymorphic; one study found 85 variants in a sample of 236 men. A small percentage of women may have an extra type of color receptor because they have different alleles for the gene for the L opsin on each X chromosome. X chromosome inactivation means that while only one opsin is expressed in each cone cell, both types may occur overall, and some women may therefore show a degree of
tetrachromat 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 te ...
ic color vision. Variations in ''
OPN1MW Green-sensitive opsin is a protein that in humans is encoded by the ''OPN1MW'' gene. OPN1MW2 is a similar opsin. See also * Opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, ...
'', which encodes the opsin expressed in M cones, appear to be rare, and the observed variants have no effect on
spectral sensitivity Spectral sensitivity is the relative efficiency of detection, of light or other signal, as a function of the frequency or wavelength of the signal. In visual neuroscience, spectral sensitivity is used to describe the different characterist ...
.


Color in the primate brain

Color processing begins at a very early level in the visual system (even within the retina) through initial color opponent mechanisms. Both Helmholtz's trichromatic theory and Hering's opponent-process theory are therefore correct, but trichromacy arises at the level of the receptors, and opponent processes arise at the level of
retinal ganglion cell A retinal ganglion cell (RGC) is a type of neuron located near the inner surface (the ganglion cell layer) of the retina of the eye. It receives visual information from photoreceptors via two intermediate neuron types: bipolar cells and reti ...
s and beyond. In Hering's theory, opponent mechanisms refer to the opposing color effect of red-green, blue-yellow, and light-dark. However, in the visual system, it is the activity of the different receptor types that are opposed. Some midget retinal ganglion cells oppose L and M cone activity, which corresponds loosely to red–green opponency, but actually runs along an axis from blue-green to magenta. Small bistratified retinal ganglion cells oppose input from the S cones to input from the L and M cones. This is often thought to correspond to blue–yellow opponency but actually runs along a color axis from yellow-green to violet. Visual information is then sent to the brain from retinal ganglion cells via the
optic nerve In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual information from the retina to the brain. In humans, the optic nerve is derived fro ...
to the
optic chiasma In neuroanatomy, the optic chiasm, or optic chiasma (; , ), is the part of the brain where the optic nerves cross. It is located at the bottom of the brain immediately inferior to the hypothalamus. The optic chiasm is found in all vertebr ...
: a point where the two optic nerves meet and information from the temporal (contralateral) visual field crosses to the other side of the brain. After the optic chiasma, the visual tracts are referred to as the optic tracts, which enter 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 direct ...
to synapse at the lateral geniculate nucleus (LGN). The lateral geniculate nucleus is divided into laminae (zones), of which there are three types: the M-laminae, consisting primarily of M-cells, the P-laminae, consisting primarily of P-cells, and the koniocellular laminae. M- and P-cells receive relatively balanced input from both L- and M-cones throughout most of the retina, although this seems to not be the case at the fovea, with midget cells synapsing in the P-laminae. The koniocellular laminae receives axons from the small bistratified ganglion cells. After synapsing at the LGN, the visual tract continues on back to the primary
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 ...
(V1) located at the back of the brain within the occipital lobe. Within V1 there is a distinct band (striation). This is also referred to as "striate cortex", with other cortical visual regions referred to collectively as "extrastriate cortex". It is at this stage that color processing becomes much more complicated. In V1 the simple three-color segregation begins to break down. Many cells in V1 respond to some parts of the spectrum better than others, but this "color tuning" is often different depending on the adaptation state of the visual system. A given cell that might respond best to long-wavelength light if the light is relatively bright might then become responsive to all wavelengths if the stimulus is relatively dim. Because the color tuning of these cells is not stable, some believe that a different, relatively small, population of neurons in V1 is responsible for color vision. These specialized "color cells" often have receptive fields that can compute local cone ratios. Such "double-opponent" cells were initially described in the goldfish retina by Nigel Daw; their existence in primates was suggested by David H. Hubel and Torsten Wiesel, first demonstrated by C.R. Michael and subsequently proven by
Bevil Conway Bevil Conway (born 4 November 1974, in Harare, Zimbabwe) is a neuroscientist and artist, and an expert in color. Conway specialises in visual perception in his scientific work, and he often explores the limitations of the visual system in his a ...
. As Margaret Livingstone and David Hubel showed, double opponent cells are clustered within localized regions of V1 called blobs, and are thought to come in two flavors, red–green and blue-yellow. Red-green cells compare the relative amounts of red-green in one part of a scene with the amount of red-green in an adjacent part of the scene, responding best to local color contrast (red next to green). Modeling studies have shown that double-opponent cells are ideal candidates for the neural machinery of color constancy explained by
Edwin H. Land Edwin Herbert Land, ForMemRS, FRPS, Hon.MRI (May 7, 1909 – March 1, 1991) was an Russian-American scientist and inventor, best known as the co-founder of the Polaroid Corporation. He invented inexpensive filters for polarizing light, ...
in his
retinex Color constancy is an example of subjective constancy and a feature of the human color perception system which ensures that the perceived color of objects remains relatively constant under varying illumination conditions. A green apple ...
theory. From the V1 blobs, color information is sent to cells in the second visual area, V2. The cells in V2 that are most strongly color tuned are clustered in the "thin stripes" that, like the blobs in V1, stain for the enzyme cytochrome oxidase (separating the thin stripes are interstripes and thick stripes, which seem to be concerned with other visual information like motion and high-resolution form). Neurons in V2 then synapse onto cells in the extended V4. This area includes not only V4, but two other areas in the posterior inferior temporal cortex, anterior to area V3, the dorsal posterior inferior temporal cortex, and posterior TEO. Area V4 was initially suggested by
Semir Zeki Semir Zeki FMedSci FRS is a British and French neurobiologist who has specialised in studying the primate visual brain and more recently the neural correlates of affective states, such as the experience of love, desire and beauty that are gen ...
to be exclusively dedicated to color, and he later showed that V4 can be subdivided into subregions with very high concentrations of color cells separated from each other by zones with lower concentration of such cells though even the latter cells respond better to some wavelengths than to others, a finding confirmed by subsequent studies. The presence in V4 of orientation-selective cells led to the view that V4 is involved in processing both color and form associated with color but it is worth noting that the orientation selective cells within V4 are more broadly tuned than their counterparts in V1, V2 and V3. Color processing in the extended V4 occurs in millimeter-sized color modules called globs. This is the part of the brain in which color is first processed into the full range of hues found in
color space A color space is a specific organization of colors. In combination with color profiling supported by various physical devices, it supports reproducible representations of colorwhether such representation entails an analog or a digital represen ...
. Anatomical studies have shown that neurons in extended V4 provide input to the inferior
temporal lobe The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe is involved i ...
. "IT" cortex is thought to integrate color information with shape and form, although it has been difficult to define the appropriate criteria for this claim. Despite this murkiness, it has been useful to characterize this pathway (V1 > V2 > V4 > IT) as the ventral stream or the "what pathway", distinguished from the dorsal stream ("where pathway") that is thought to analyze motion, among other features.


Subjectivity of color perception

Color is a feature of visual perception by an observer. There is a complex relationship between the wavelengths of light in the visual spectrum and human experiences of color. Although most people are assumed to have the same mapping, the philosopher
John Locke John Locke (; 29 August 1632 – 28 October 1704) was an English philosopher and physician, widely regarded as one of the most influential of Enlightenment thinkers and commonly known as the "father of liberalism". Considered one of ...
recognized that alternatives are possible, and described one such hypothetical case with the " inverted spectrum" thought experiment. For example, someone with an inverted spectrum might experience green while seeing 'red' (700 nm) light, and experience red while seeing 'green' (530 nm) light. This inversion has never been demonstrated in experiment, though.
Synesthesia Synesthesia (American English) or synaesthesia (British English) is a perceptual phenomenon in which stimulation of one sensory or cognitive pathway leads to involuntary experiences in a second sensory or cognitive pathway. People who re ...
(or
ideasthesia Ideasthesia (alternative spelling ideaesthesia) is a neuropsychological phenomenon in which activations of concepts (inducers) evoke perception-like sensory experiences (concurrents). The name comes from the Ancient Greek () and (), meaning 'se ...
) provides some atypical but illuminating examples of subjective color experience triggered by input that is not even light, such as sounds or shapes. The possibility of a clean dissociation between color experience from properties of the world reveals that color is a subjective psychological phenomenon. The Himba people have been found to categorize colors differently from most Westerners and are able to easily distinguish close shades of green, barely discernible for most people. The Himba have created a very different color scheme which divides the spectrum to dark shades (''zuzu'' in Himba), very light (''vapa''), vivid blue and green (''buru'') and dry colors as an adaptation to their specific way of life. The perception of color depends heavily on the context in which the perceived object is presented. Psychophysical experiments have shown that color is perceived before the orientation of lines and directional motion by as much as 40ms and 80 ms respectively, thus leading to a perceptual asynchrony that is demonstrable with brief presentation times.


Chromatic adaptation

In color vision, chromatic adaptation refers to color constancy; the ability of the visual system to preserve the appearance of an object under a wide range of light sources. For example, a white page under blue, pink, or purple light will reflect mostly blue, pink, or purple light to the eye, respectively; the brain, however, compensates for the effect of lighting (based on the color shift of surrounding objects) and is more likely to interpret the page as white under all three conditions, a phenomenon known as color constancy. In color science, chromatic adaptation is the estimation of the representation of an object under a different light source from the one in which it was recorded. A common application is to find a ''chromatic adaptation transform'' (CAT) that will make the recording of a neutral object appear neutral (
color balance In photography and image processing, color balance is the global adjustment of the intensities of the colors (typically red, green, and blue primary colors). An important goal of this adjustment is to render specific colors – particularly ne ...
), while keeping other colors also looking realistic. For example, chromatic adaptation transforms are used when converting images between ICC profiles with different white points.
Adobe Photoshop Adobe Photoshop is a raster graphics editor developed and published by Adobe Inc. for Windows and macOS. It was originally created in 1988 by Thomas and John Knoll. Since then, the software has become the industry standard not only in rast ...
, for example, uses the Bradford CAT.


Color vision in nonhumans

Many species can see light with frequencies outside the human "
visible spectrum The visible spectrum is the portion of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called '' visible light'' or simply light. A typical human eye will respond to ...
".
Bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s and many other insects can detect ultraviolet light, which helps them to find nectar in flowers. Plant species that depend on insect pollination may owe reproductive success to ultraviolet "colors" and patterns rather than how colorful they appear to humans. Birds, too, can see into the ultraviolet (300–400 nm), and some have sex-dependent markings on their plumage that are visible only in the ultraviolet range. Many animals that can see into the ultraviolet range, however, cannot see red light or any other reddish wavelengths. For example, bees' visible spectrum ends at about 590 nm, just before the orange wavelengths start. Birds, however, can see some red wavelengths, although not as far into the light spectrum as humans. It is a myth that the common goldfish is the only animal that can see both infrared and ultraviolet light; their color vision extends into the ultraviolet but not the infrared. The basis for this variation is the number of cone types that differ between species. Mammals, in general, have a color vision of a limited type, and usually have
red-green color blindness Color blindness or color vision deficiency (CVD) is the decreased ability to see color or differences in color. It can impair tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights. Color blindness may make some aca ...
, with only two types of cones. Humans, some primates, and some marsupials see an extended range of colors, but only by comparison with other mammals. Most non-mammalian vertebrate species distinguish different colors at least as well as humans, and many species of birds, fish, reptiles, and amphibians, and some invertebrates, have more than three cone types and probably superior color vision to humans. In most Catarrhini (Old World monkeys and apes—primates closely related to humans), there are three types of
color receptors 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. Co ...
(known as
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. Cone ...
s), resulting in trichromatic color vision. These primates, like humans, are known as
trichromats Trichromacy or trichromatism is the possessing of three independent channels for conveying color information, derived from the three different types of cone cells in the eye. Organisms with trichromacy are called trichromats. The normal expl ...
. Many other primates (including New World monkeys) and other mammals are
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 requir ...
s, which is the general color vision state for mammals that are active during the day (i.e., felines, canines, ungulates). Nocturnal mammals may have little or no color vision. Trichromat non-primate mammals are rare. Many
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s have color vision. Honeybees and
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s have trichromatic color vision which is insensitive to red but sensitive to ultraviolet. '' Osmia rufa'', for example, possess a trichromatic color system, which they use in foraging for pollen from flowers. In view of the importance of color vision to bees one might expect these receptor sensitivities to reflect their specific visual ecology; for example the types of flowers that they visit. However, the main groups of hymenopteran insects excluding ants (i.e., bees, wasps and sawflies) mostly have three types of photoreceptor, with spectral sensitivities similar to the honeybee's.'' Papilio'' butterflies possess six types of photoreceptors and may have pentachromatic vision. The most complex color vision system in the animal kingdom has been found in stomatopods (such as the mantis shrimp) having between 12 and 16 spectral receptor types thought to work as multiple dichromatic units. Vertebrate animals such as tropical fish and birds sometimes have more complex color vision systems than humans; thus the many subtle colors they exhibit generally serve as direct signals for other fish or birds, and not to signal mammals. In bird vision,
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 te ...
is achieved through up to four cone types, depending on species. Each single cone contains one of the four main types of vertebrate cone photopigment (LWS/ MWS, RH2, SWS2 and SWS1) and has a colored oil droplet in its inner segment. Brightly colored oil droplets inside the cones shift or narrow the spectral sensitivity of the cell. Pigeons may be
pentachromat 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 ...
s. Reptiles and amphibians also have four cone types (occasionally five), and probably see at least the same number of colors that humans do, or perhaps more. In addition, some nocturnal geckos and
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s have the capability of seeing color in dim light. At least some color-guided behaviors in amphibians have also been shown to be wholly innate, developing even in visually deprived animals. In the
evolution of mammals The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked l ...
, segments of color vision were lost, then for a few species of primates, regained by gene duplication. Eutherian mammals other than primates (for example, dogs, mammalian farm animals) generally have less-effective two-receptor (
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 requir ...
ic) color perception systems, which distinguish blue, green, and yellow—but cannot distinguish oranges and reds. There is some evidence that a few mammals, such as cats, have redeveloped the ability to distinguish longer wavelength colors, in at least a limited way, via one-amino-acid mutations in opsin genes. The adaptation to see reds is particularly important for primate mammals, since it leads to the identification of fruits, and also newly sprouting reddish leaves, which are particularly nutritious. However, even among primates, full color vision differs between New World and Old World monkeys. Old World primates, including monkeys and all apes, have vision similar to humans. New World monkeys may or may not have color sensitivity at this level: in most species, males are dichromats, and about 60% of females are trichromats, but the owl monkeys are cone
monochromats 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 ...
, and both sexes of howler monkeys are trichromats. Visual sensitivity differences between males and females in a single species is due to the gene for yellow-green sensitive
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 ...
protein (which confers ability to differentiate red from green) residing on the X sex chromosome. Several
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 ...
s, such as the
fat-tailed dunnart The fat-tailed dunnart (''Sminthopsis crassicaudata'') is a species of mouse-like marsupial of the Dasyuridae, the family that includes the little red kaluta, quolls, and the Tasmanian devil. It has an average body length of with a tail of . Ea ...
(''Sminthopsis crassicaudata''), have trichromatic color vision.
Marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...
s, adapted for low-light vision, have only a single cone type and are thus monochromats.


Evolution

Color perception mechanisms are highly dependent on evolutionary factors, of which the most prominent is thought to be satisfactory recognition of food sources. In herbivorous primates, color perception is essential for finding proper (immature) leaves. In hummingbirds, particular flower types are often recognized by color as well. On the other hand,
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
mammals have less-developed color vision since adequate light is needed for cones to function properly. There is evidence that
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 radiation ...
light plays a part in color perception in many branches of the animal kingdom, especially
insect 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 pa ...
s. In general, the optical spectrum encompasses the most common electronic transitions in the matter and is therefore the most useful for collecting information about the environment. The evolution of trichromatic color vision in primates occurred as the ancestors of modern monkeys, apes, and humans switched to diurnal (daytime) activity and began consuming fruits and leaves from flowering plants. Color vision, with UV discrimination, is also present in a number of arthropods—the only terrestrial animals besides the vertebrates to possess this trait. Some animals can distinguish colors in the ultraviolet spectrum. The UV spectrum falls outside the human visible range, except for some cataract surgery patients. Birds, turtles, lizards, many fish and some rodents have UV receptors in their retinas. These animals can see the UV patterns found on flowers and other wildlife that are otherwise invisible to the human eye. Ultraviolet vision is an especially important adaptation in birds. It allows birds to spot small prey from a distance, navigate, avoid predators, and forage while flying at high speeds. Birds also utilize their broad spectrum vision to recognize other birds, and in sexual selection.


Mathematics of color perception

A "physical color" is a combination of pure spectral colors (in the visible range). In principle there exist infinitely many distinct spectral colors, and so the set of all physical colors may be thought of as an infinite-dimensional
vector space In mathematics and physics, a vector space (also called a linear space) is a set whose elements, often called '' vectors'', may be added together and multiplied ("scaled") by numbers called ''scalars''. Scalars are often real numbers, but can ...
(a
Hilbert space In mathematics, Hilbert spaces (named after David Hilbert) allow generalizing the methods of linear algebra and calculus from (finite-dimensional) Euclidean vector spaces to spaces that may be infinite-dimensional. Hilbert spaces arise natural ...
). This space is typically notated ''H''color. More technically, the space of physical colors may be considered to be the topological cone over the
simplex In geometry, a simplex (plural: simplexes or simplices) is a generalization of the notion of a triangle or tetrahedron to arbitrary dimensions. The simplex is so-named because it represents the simplest possible polytope in any given dimension. ...
whose vertices are the spectral colors, with white at the centroid of the simplex, black at the apex of the cone, and the monochromatic color associated with any given vertex somewhere along the line from that vertex to the apex depending on its brightness. An element ''C'' of ''H''color is a function from the range of visible wavelengths—considered as an interval of real numbers 'W''min,''W''max��to the real numbers, assigning to each wavelength ''w'' in 'W''min,''W''maxits intensity ''C''(''w''). A humanly perceived color may be modeled as three numbers: the extents to which each of the 3 types of cones is stimulated. Thus a humanly perceived color may be thought of as a point in 3-dimensional
Euclidean space Euclidean space is the fundamental space of geometry, intended to represent physical space. Originally, that is, in Euclid's ''Elements'', it was the three-dimensional space of Euclidean geometry, but in modern mathematics there are Euclidea ...
. We call this space R3color. Since each wavelength ''w'' stimulates each of the 3 types of cone cells to a known extent, these extents may be represented by 3 functions ''s''(''w''), ''m''(''w''), ''l''(''w'') corresponding to the response of the ''S'', ''M'', and ''L'' cone cells, respectively. Finally, since a beam of light can be composed of many different wavelengths, to determine the extent to which a physical color ''C'' in ''H''color stimulates each cone cell, we must calculate the integral (with respect to ''w''), over the interval 'W''min,''W''max of ''C''(''w'')·''s''(''w''), of ''C''(''w'')·''m''(''w''), and of ''C''(''w'')·''l''(''w''). The triple of resulting numbers associates with each physical color ''C'' (which is an element in ''H''color) a particular perceived color (which is a single point in R3color). This association is easily seen to be linear. It may also easily be seen that many different elements in the "physical" space ''H''color can all result in the same single perceived color in R3color, so a perceived color is not unique to one physical color. Thus human color perception is determined by a specific, non-unique linear mapping from the infinite-dimensional Hilbert space ''H''color to the 3-dimensional Euclidean space R3color. Technically, the image of the (mathematical) cone over the simplex whose vertices are the spectral colors, by this linear mapping, is also a (mathematical) cone in R3color. Moving directly away from the vertex of this cone represents maintaining the same chromaticity while increasing its intensity. Taking a cross-section of this cone yields a 2D chromaticity space. Both the 3D cone and its projection or cross-section are convex sets; that is, any mixture of spectral colors is also a color. In practice, it would be quite difficult to physiologically measure an individual's three cone responses to various physical color stimuli. Instead, a psychophysical approach is taken. Three specific benchmark test lights are typically used; let us call them ''S'', ''M'', and ''L''. To calibrate human perceptual space, scientists allowed human subjects to try to match any physical color by turning dials to create specific combinations of intensities (''I''''S'', ''I''''M'', ''I''''L'') for the ''S'', ''M'', and ''L'' lights, resp., until a match was found. This needed only to be done for physical colors that are spectral, since a linear combination of spectral colors will be matched by the same linear combination of their (''I''''S'', ''I''''M'', ''I''''L'') matches. Note that in practice, often at least one of ''S'', ''M'', ''L'' would have to be added with some intensity to the ''physical test color'', and that combination matched by a linear combination of the remaining 2 lights. Across different individuals (without color blindness), the matchings turned out to be nearly identical. By considering all the resulting combinations of intensities (''I''''S'', ''I''''M'', ''I''''L'') as a subset of 3-space, a model for human perceptual color space is formed. (Note that when one of ''S'', ''M'', ''L'' had to be added to the test color, its intensity was counted as negative.) Again, this turns out to be a (mathematical) cone, not a quadric, but rather all rays through the origin in 3-space passing through a certain convex set. Again, this cone has the property that moving directly away from the origin corresponds to increasing the intensity of the ''S'', ''M'', ''L'' lights proportionately. Again, a cross-section of this cone is a planar shape that is (by definition) the space of "chromaticities" (informally: distinct colors); one particular such cross-section, corresponding to constant ''X''+''Y''+''Z'' of the CIE 1931 color space, gives the CIE chromaticity diagram. This system implies that for any hue or non-spectral color not on the boundary of the chromaticity diagram, there are infinitely many distinct physical spectra that are all perceived as that hue or color. So, in general, there is no such thing as ''the'' combination of spectral colors that we perceive as (say) a specific version of tan; instead, there are infinitely many possibilities that produce that exact color. The boundary colors that are pure spectral colors can be perceived only in response to light that is purely at the associated wavelength, while the boundary colors on the "line of purples" can each only be generated by a specific ratio of the pure violet and the pure red at the ends of the visible spectral colors. The CIE chromaticity diagram is horseshoe-shaped, with its curved edge corresponding to all spectral colors (the ''spectral locus''), and the remaining straight edge corresponding to the most saturated
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 ...
s, mixtures of red and violet.


See also

* Achromatopsia * Color blindness * Color theory * Inverted spectrum * Primary color *
The dress The dress was a viral phenomenon on the Internet in 2015. Viewers of the image disagreed on whether the dress depicted in a photograph was coloured black and blue, or white and gold. The phenomenon revealed differences in human colour percept ...
*
Visual perception Visual perception is the ability to interpret the surrounding environment through photopic vision (daytime vision), color vision, scotopic vision (night vision), and mesopic vision (twilight vision), using light in the visible spectrum ref ...


References


Further reading

* * * * * {{Authority control Image processing Visual perception Mathematics in medicine