Color vision

Color vision is the ability of an organism or machine to distinguish objects based on the wavelengths (or frequencies) of the light they reflect, emit, or transmit. Colors can be measured and quantified in various ways; indeed, a person's perception of colors is a subjective process whereby the brain responds to the stimuli that are produced when incoming light reacts with the several types of cone cells in the eye. In essence, different people see the same illuminated object or light source in different ways.

Wavelength and hue detection

Isaac Newton discovered that white light, after being split into its component colours when passed through a dispersive prism, could be recombined to make white light by passing them through a different prism.

The characteristic colours are, from long to short wavelengths (and, correspondingly, from low to high frequency), red, orange, yellow, green, blue, and violet. 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 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. Rods are maximally sensitive to wavelengths near 500 nm, and play little, if any, role in colour vision. In brighter light, such as daylight, vision is photopic: light is detected by cone cells which are responsible for colour 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 cells. The shift in colour 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 colours of just a few wavelengths in animals with few types of colour receptors. In humans, white light can be perceived by combining wavelengths such as red, green, and blue, or just a pair of complementary colours such as blue and yellow.[1]

Physiology of color perception

Modern Color Vision Model
The modern model of human color perception as it occurs in the retina, pertaining to both the trichromatic and opponent process theories introduced in the 19th century.
Cone-fundamentals-with-srgb-spectrum
Normalized response spectra of human cones, to monochromatic spectral stimuli, with wavelength given in nanometers.
Spectrum locus 12
The same figures as above represented here as a single curve in three (normalized cone response) dimensions
Eyesensitivity
Photopic relative brightness sensitivity of the human visual system as a function of wavelength (luminosity function)

Perception of color begins with specialized retinal cells containing pigments with different spectral sensitivities, known as cone cells. In humans, there are three types of cones sensitive to three different spectra, resulting in trichromatic color vision.

Each individual cone contains pigments composed of opsin apoprotein, which is covalently linked to either 11-cis-hydroretinal or more rarely 11-cis-dehydroretinal.[2]

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 it will be finalized in the visual cortex 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- and M-cones do not directly correspond to blue and green, although they are often described as such. The RGB color model, 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;[3] in some non-human species this polymorphic variation is even greater, and it may well be adaptive.[4]

Another sensitivity maximum of the red-sensitive cones

Some sources (e.g.[5]) assert that the erythropsin in the red-sensitive cones has two ranges of sensitivities. The main (red) range has its maximum at 600 nm. The tiny (violet) range is between 380 nm and 450 nm, peaking at 420 nm. This small maximum is said to explain why the visible light with the shortest wavelength looks violet (rather than dark blue), while the visible light with the longest wavelength looks dark red.

Theories

Two complementary theories of color vision are the trichromatic theory and the opponent process theory. The trichromatic theory, or Young–Helmholtz theory, proposed in the 19th century by Thomas Young and Hermann von Helmholtz, as mentioned above, states that the retina's three types of cones are preferentially sensitive to blue, green, and red. Ewald Hering proposed the opponent process theory in 1872.[6] It states that the visual system interprets color in an antagonistic way: red vs. green, blue vs. yellow, black vs. white. Both theories are now accepted as valid, describing different stages in visual physiology, visualized in the adjacent diagram.[7] 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.

Cone cells in the human eye

Cone type Name Range Peak wavelength[8][9]
S β 400–500 nm 420–440 nm
M γ 450–630 nm 534–555 nm
L ρ 500–700 nm 564–580 nm

A range of wavelengths of light stimulates each of these receptor types to varying degrees. Yellowish-green light, for example, stimulates both L and M cones equally strongly, but only stimulates S-cones weakly. Red light, on the other hand, stimulates L cones much more than M cones, and S cones hardly at all; blue-green light stimulates M cones more than L cones, and S cones a bit more strongly, and is also the peak stimulant for rod cells; and blue light stimulates S cones more strongly than red or green light, but L and M cones more weakly. The brain combines the information from each type of receptor to give rise to different perceptions of different wavelengths of light.

The opsins (photopigments) present in the L and M cones are encoded on the X chromosome; defective encoding of these leads to the two most common forms of color blindness. The OPN1LW gene, which codes for the opsin present in the L cones, is highly polymorphic (a recent study by Verrelli and Tishkoff found 85 variants in a sample of 236 men).[10] A very 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 occur overall, and some women may therefore show a degree of tetrachromatic color vision.[11] Variations in OPN1MW, which codes the opsin expressed in M cones, appear to be rare, and the observed variants have no effect on spectral sensitivity.

Color in the human brain

Ventral-dorsal streams
Visual pathways in the human brain. The ventral stream (purple) is important in color recognition. The dorsal stream (green) is also shown. They originate from a common source in the visual cortex.

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 cells 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 to the optic chiasma: 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 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 receive axons from the small bistratified ganglion cells.[12][13]

After synapsing at the LGN, the visual tract continues on back to the primary visual cortex (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;[14][15] their existence in primates was suggested by David H. Hubel and Torsten Wiesel and subsequently proven by Bevil Conway.[16] 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.[17] 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 in his retinex theory.[18]

16777216colors
When viewed in full size, this image contains about 16 million pixels, each corresponding to a different color on the full set of RGB colors. The human eye can distinguish about 10 million different colors.[19]

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.[20][21] Area V4 was initially suggested by Semir Zeki to be exclusively dedicated to color, but this is now thought to be incorrect.[22] In particular, 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.[23] Color processing in the extended V4 occurs in millimeter-sized color modules called globs.[20][21] This is the first part of the brain in which color is processed in terms of the full range of hues found in color space.[20][21]

Anatomical studies have shown that neurons in extended V4 provide input to the inferior temporal lobe . "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 many other features.

Subjectivity of color perception

Nothing categorically distinguishes the visible spectrum of electromagnetic radiation from invisible portions of the broader spectrum. In this sense, color is not a property of electromagnetic radiation, but a feature of visual perception by an observer. Furthermore, there is an arbitrary mapping between 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 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. Synesthesia (or ideasthesia) 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 Euro-Americans and are able to easily distinguish close shades of green, barely discernible for most people.[24] 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.

Perception of color depends heavily on the context in which the perceived object is presented. 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 other animal species

Many species can see light with frequencies outside the human "visible spectrum". Bees 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.[25][26] 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.[27] It is an incorrect popular belief that the common goldfish is the only animal that can see both infrared and ultraviolet light;[28] their color vision extends into the ultraviolet but not the infrared.[29]

The basis for this variation is the number of cone types that differ between species. Mammals in general have color vision of a limited type, and usually have red-green color blindness, 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 (known as cone cells), resulting in trichromatic color vision. These primates, like humans, are known as trichromats. Many other primates (including New World monkeys) and other mammals are dichromats, 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.[30][31]

Many invertebrates have color vision. Honeybees and bumblebees 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.[32] 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.[33] Papilio butterflies possess six types of photoreceptors and may have pentachromatic vision.[34] The most complex color vision system in the animal kingdom has been found in stomatopods (such as the mantis shrimp) with up to 12 spectral receptor types thought to work as multiple dichromatic units.[35]

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.[36] In bird vision, tetrachromacy 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.[33] Brightly colored oil droplets inside the cones shift or narrow the spectral sensitivity of the cell. It has been suggested that it is likely that pigeons are pentachromats.[37]

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 have the capability of seeing color in dim light.[38]

In the evolution of mammals, 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 (dichromatic) 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.[39] The adaptation to see reds is particularly important for primate mammals, since it leads to 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, and both sexes of howler monkeys are trichromats.[40][41][42][43] Visual sensitivity differences between males and females in a single species is due to the gene for yellow-green sensitive opsin protein (which confers ability to differentiate red from green) residing on the X sex chromosome.

Several marsupials such as the fat-tailed dunnart (Sminthopsis crassicaudata) have been shown to have trichromatic color vision.[44]

Marine mammals, adapted for low-light vision, have only a single cone type and are thus monochromats.

Color vision table
State Types of cone cells Approx. number of colors perceived Carriers
Monochromacy 1 200 Marine mammals, owl monkey, Australian sea lion, achromat primates
Dichromacy 2 40,000 Most terrestrial non-primate mammals, color blind primates
Trichromacy 3 10 million[45] Most primates, especially great apes (such as humans), marsupials, some insects (such as honeybees)
Tetrachromacy 4 100 million Most reptiles, amphibians, birds and insects, rarely humans
Pentachromacy 5 10 billion Some insects (specific species of butterflies), some birds (pigeons for instance)

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 mammals have less-developed color vision, since adequate light is needed for cones to function properly. There is evidence that ultraviolet light plays a part in color perception in many branches of the animal kingdom, especially insects. In general, the optical spectrum encompasses the most common electronic transitions in 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.[46] Color vision, with UV discrimination, is also present in a number of arthropods—the only terrestrial animals besides the vertebrates to possess this trait.[47]

Some animals can distinguish colors in the ultraviolet spectrum. The UV spectrum falls outside the human visible range, except for some cataract surgery patients.[48] Birds, turtles, lizards, many fish and some rodents have UV receptors in their retinas.[49] 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.[50][51]

Mathematics of color perception

A "physical color" is a combination of pure spectral colors (in the visible range). Since there are, in principle, infinitely many distinct spectral colors, the set of all physical colors may be thought of as an infinite-dimensional vector space, in fact a Hilbert space. We call this space Hcolor. More technically, the space of physical colors may be considered to be the (mathematical) cone over the simplex 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 Hcolor is a function from the range of visible wavelengths—considered as an interval of real numbers [Wmin,Wmax]—to the real numbers, assigning to each wavelength w in [Wmin,Wmax] its 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. 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 Hcolor stimulates each cone cell, we must calculate the integral (with respect to w), over the interval [Wmin,Wmax], of C(ws(w), of C(wm(w), and of C(wl(w). The triple of resulting numbers associates with each physical color C (which is an element in Hcolor) 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 Hcolor 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 Hcolor 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.

Cie Chart with sRGB gamut by spigget
The CIE 1931 xy chromaticity diagram with a triangle showing the gamut of the sRGB color space. The Planckian locus is shown with color temperatures labeled in kelvins. The outer curved boundary is the spectral (or monochromatic) locus, with wavelengths shown in nanometers (blue). Note that the colors in this file are being specified using sRGB. Areas outside the triangle cannot be accurately rendered because they are out of the gamut of sRGB, therefore they have been interpreted. Note that the colors depicted depend on the color space of the device you use to view the image (number of colors on your monitor, etc.), and may not be a strictly accurate representation of the color at a particular position.

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 (IS, IM, IL) 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 (IS, IM, IL) 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 (IS, IM, IL) 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 purples, mixtures of red and violet.

Chromatic adaptation

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), while keeping other colors also looking realistic.[52] For example, chromatic adaptation transforms are used when converting images between ICC profiles with different white points. Adobe Photoshop, for example, uses the Bradford CAT.[53]

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.[54]

See also

References

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External links

Achromatopsia

Achromatopsia (ACHM), also known as total color blindness, is a medical syndrome that exhibits symptoms relating to at least five conditions. The term may refer to acquired conditions such as cerebral achromatopsia, but it typically refers to an autosomal recessive congenital color vision condition, the inability to perceive color and to achieve satisfactory visual acuity at high light levels (typically exterior daylight). The syndrome is also present in an incomplete form which is more properly defined as dyschromatopsia. It is estimated to affect 1 in 30,000 live births worldwide.

There is some discussion as to whether achromats can see color or not. As illustrated in The Island of the Colorblind by Oliver Sacks, some achromats cannot see color, only black, white, and shades of grey. With five different genes currently known to cause similar symptoms, it may be that some do see marginal levels of color differentiation due to different gene characteristics. With such small sample sizes and low response rates, it is difficult to accurately diagnose the 'typical achromatic conditions'. If the light level during testing is optimized for them, they may achieve corrected visual acuity of 20/100 to 20/150 at lower light levels, regardless of the absence of color.

One common trait is hemeralopia or blindness in full sun. In patients with achromatopsia, the cone system and fibres carrying color information remain intact. This indicates that the mechanism used to construct colors is defective.

Bee learning and communication

Honey bees are sensitive to odors (including pheromones), tastes, and colors, including ultraviolet. They can demonstrate capabilities such as color discrimination through classical and operant conditioning and retain this information for several days at least; they communicate the location and nature of sources of food; they adjust their foraging to the times at which food is available; they may even form cognitive maps of their surroundings.

Color

Color (American English), or colour (Commonwealth English), is the characteristic of human visual perception described through color categories, with names such as red, orange, yellow, green, blue, or purple. This perception of color derives from the stimulation of cone cells in the human eye by electromagnetic radiation in the visible spectrum. Color categories and physical specifications of color are associated with objects through the wavelength of the light that is reflected from them. This reflection is governed by the object's physical properties such as light absorption, emission spectra, etc.

By defining a color space, colors can be identified numerically by coordinates, which in 1931 were also named in global agreement with internationally agreed color names like mentioned above (red, orange, etc.) by the International Commission on Illumination. The RGB color space for instance is a color space corresponding to human trichromacy and to the three cone cell types that respond to three bands of light: long wavelengths, peaking near 564–580 nm (red); medium-wavelength, peaking near 534–545 nm (green); and short-wavelength light, near 420–440 nm (blue). There may also be more than three color dimensions in other color spaces, such as in the CMYK color model, wherein one of the dimensions relates to a color's colorfulness).

The photo-receptivity of the "eyes" of other species also varies considerably from that of humans and so results in correspondingly different color perceptions that cannot readily be compared to one another. Honeybees and bumblebees for instance have trichromatic color vision sensitive to ultraviolet but is insensitive to red. 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) with up to 12 spectral receptor types thought to work as multiple dichromatic units.The science of color is sometimes called chromatics, colorimetry, or simply color science. It includes the study of the perception of color by the human eye and brain, the origin of color in materials, color theory in art, and the physics of electromagnetic radiation in the visible range (that is, what is commonly referred to simply as light).

Color blindness

Color blindness, also known as color vision deficiency, is the decreased ability to see color or differences in color. Simple tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights can be more challenging. Color blindness may also make some educational activities more difficult. However, problems are generally minor, and most people find that they can adapt. People with total color blindness (achromatopsia) may also have decreased visual acuity and be uncomfortable in bright environments.The most common cause of color blindness is an inherited problem in the development of one or more of the three sets of color sensing cones in the eye. Males are more likely to be color blind than females, as the genes responsible for the most common forms of color blindness are on the X chromosome. As females have two X chromosomes, a defect in one is typically compensated for by the other, while males only have one X chromosome. Color blindness can also result from physical or chemical damage to the eye, optic nerve or parts of the brain. Diagnosis is typically with the Ishihara color test; however, a number of other testing methods also exist.There is no cure for color blindness. Diagnosis may allow a person's teacher to change their method of teaching to accommodate the decreased ability to recognize colors. Special lenses may help people with red-green color blindness when under bright conditions. There are also mobile apps that can help people identify colors.Red-green color blindness is the most common form, followed by blue-yellow color blindness and total color blindness. Red-green color blindness affects up to 8% of males and 0.5% of females of Northern European descent. The ability to see color also decreases in old age. Being color blind may make people ineligible for certain jobs in certain countries. This may include being a pilot, train driver and working in the armed forces. The effect of color blindness on artistic ability, however, is controversial. The ability to draw appears to be unchanged, and a number of famous artists are believed to have been color blind.

Color constancy

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 for instance looks green to us at midday, when the main illumination is white sunlight, and also at sunset, when the main illumination is red. This helps us identify objects.

Color model

A color model is an abstract mathematical model describing the way colors can be represented as tuples of numbers, typically as three or four values or color components. When this model is associated with a precise description of how the components are to be interpreted (viewing conditions, etc.), the resulting set of colors is called "color space." This section describes ways in which human color vision can be modeled.

Dichromacy

Dichromacy is the state of having two types of functioning color receptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats can match any color they see with a mixture of no more than two pure spectral lights. By comparison, trichromats require three pure spectral lights to match all colors that they can perceive, and tetrachromats require four.

Dichromacy in humans is a color vision defect in which one of the three basic color mechanisms is absent or not functioning. It is hereditary and sex-linked, predominantly affecting males. Dichromacy occurs when one of the cone pigments is missing and color is reduced to two dimensions. The term is from di meaning "two" and chroma meaning "color".

Dichromatism

Dichromatism (or polychromatism) is a phenomenon where a material or solution's hue is dependent on both the concentration of the absorbing substance and the depth or thickness of the medium traversed. In most substances which are not dichromatic, only the brightness and saturation of the colour depend on their concentration and layer thickness.

Examples of dichromatic substances are pumpkin seed oil, bromophenol blue, and resazurin.

When the layer of pumpkin seed oil is less than 0.7 mm thick, the oil appears bright green, and in layer thicker than this, it appears bright red.

The phenomenon is related to both the physical chemistry properties of the substance and the physiological response of the human visual system to colour. This combined physicochemical–physiological basis was first explained in 2007.

Evolution of color vision

Color vision, a proximate adaptation of the vision sensory modality, allows for the discrimination of light based on its wavelength components.

Evolution of color vision in primates

The evolution of color vision in primates is unique compared to most eutherian mammals. A remote vertebrate ancestor of primates possessed tetrachromacy, but nocturnal, warm-blooded, mammalian ancestors lost two of four cones in the retina at the time of dinosaurs. Most teleost fish, reptiles and birds are therefore tetrachromatic while most mammals are strictly dichromats, the exceptions being some primates and marsupials, who are trichromats, and many marine mammals, who are monochromats.

Primates achieve trichromacy through color photoreceptors (cone cells), with spectral peaks in the violet (short wave, S), green (middle wave, M), and yellow-green (long wave, L) wavelengths. Opsin is the primary photopigment in primate eyes, and the sequence of an organism's opsin proteins determines the spectral sensitivity of its cone cells. Not all primates, however, are capable of trichromacy. The catarrhines (Old World monkeys and apes) are routine trichromats, meaning both males and females possess three opsins (pigments) sensitive to short-, medium-, and long wavelengths. In nearly all species of platyrrhines (New World monkeys) males and homozygous females are dichromats, while heterozygous females are trichromats, a condition known as allelic or polymorphic trichromacy. Among platyrrhines, the exceptions are Alouatta (consistent trichromats) and Aotus (consistent monochromats).

Ishihara test

The Ishihara test is a color perception test for red-green color deficiencies, the first in a class of successful color vision tests called pseudo-isochromatic plates ("PIP"). It was named after its designer, Dr. Shinobu Ishihara, a professor at the University of Tokyo, who first published his tests in 1917.The test consists of a number of colored plates, called Ishihara plates, each of which contains a circle of dots appearing randomized in color and size. Within the pattern are dots which form a number or shape clearly visible to those with normal color vision, and invisible, or difficult to see, to those with a red-green color vision defect. Other plates are intentionally designed to reveal numbers only to those with a red/green color vision deficiency, and be invisible to those with normal red/green color vision. The full test consists of 38 plates, but the existence of a severe deficiency is usually apparent after only a few plates. There are also Ishihara tests consisting of 10, 14 or 24 test plates.

Monochromacy

Monochromacy (from Greek mono, meaning "one "and chromo, meaning "color") is the ability of organisms or machines to distinguish only one single frequency of the electromagnetic light spectrum. In the physical sense, no source of electromagnetic radiation is purely monochromatic but can be considered as a gaussian distribution of frequencies shaped around a peak. In the same way, a visual system of an organism or a machine cannot be monochromat but will distinguish a continuous set of frequencies around a peak, depending by the intensity of the light. Organisms with monochromacy are called monochromats.

Many species, such as all marine mammals, the owl monkey and the Australian sea lion (pictured at right) are monochromats under normal conditions. In humans, absence of color discrimination or poor color discrimination is one among several other symptoms of severe inherited or acquired diseases, as for example inherited achromatopsia, acquired achromatopsia or inherited blue cone monochromacy.

OPN1SW

Blue-sensitive opsin is a protein that in humans is encoded by the OPN1SW gene.

Opponent process

The color opponent process is a color theory that states that the human visual system interprets information about color by processing signals from cones and rods in an antagonistic manner. The three types of cones (L for long, M for medium and S for short) have some overlap in the wavelengths of light to which they respond, so it is more efficient for the visual system to record differences between the responses of cones, rather than each type of cone's individual response. The opponent color theory suggests that there are three opponent channels: red versus green, blue versus yellow, and black versus white (the last type is achromatic and detects light-dark variation, or luminance). Responses to one color of an opponent channel are antagonistic to those to the other color. That is, opposite opponent colors are never perceived together – there is no "greenish red" or "yellowish blue".

While the trichromatic theory defines the way the retina of the eye allows the visual system to detect color with three types of cones, the opponent process theory accounts for mechanisms that receive and process information from cones. Though the trichromatic and opponent processes theories were initially thought to be at odds, it later came to be understood that the mechanisms responsible for the opponent process receive signals from the three types of cones and process them at a more complex level.Besides the cones, which detect light entering the eye, the biological basis of the opponent theory involves two other types of cells: bipolar cells, and ganglion cells. Information from the cones is passed to the bipolar cells in the retina, which may be the cells in the opponent process that transform the information from cones. The information is then passed to ganglion cells, of which there are two major classes: magnocellular, or large-cell layers, and parvocellular, or small-cell layers. Parvocellular cells, or P cells, handle the majority of information about color, and fall into two groups: one that processes information about differences between firing of L and M cones, and one that processes differences between S cones and a combined signal from both L and M cones. The first subtype of cells are responsible for processing red–green differences, and the second process blue–yellow differences. P cells also transmit information about intensity of light (how much of it there is) due to their receptive fields.

Philosophy of color

Within the philosophy of color, there is a dispute between color realism, the view that colors are physical properties that objects possess, and color fictionalism, a species of error theory viewing colors according to which there are no such physical properties that objects possess.

Primary color

A set of primary colors is, most tangibly, a set of real colorants or colored lights that can be combined in varying amounts to produce a gamut of colors. This is the essential method used in applications that are intended to elicit the perception of diverse sets of color, e.g. electronic displays, color printing, and paintings. Perceptions associated with a given combination of primary colors are predicted by applying the appropriate mixing model (additive, subtractive, additive averaging etc.) that embodies the underlying physics of how light interacts with the media and ultimately the retina.

Primary colors can also be conceptual, either as additive mathematical elements of a color space or as irreducible phenomenological categories in domains such as psychology and philosophy. Color-space primaries are precisely defined and empirically rooted in psychophysical color matching experiments which are foundational for understanding color vision. Primaries of some color spaces are complete (that is, all visible colors are described in terms of their weighted sums with nonnegative weights) but necessarily imaginary (that is, there is no plausible way that those primary colors could be represented physically, or perceived). Phenomenological accounts of primary colors, such as the psychological primaries, have been used as the conceptual basis for practical color applications even though they are not a quantitative description in and of themselves.

All sets of real and color-space primaries are arbitrary, in the sense that there is no one set of primaries that can be considered the canonical set. Primary pigments or light sources selected for a given application on the basis of subjective preferences as well as practical factors such as cost, stability, availability etc. Color-space primaries can be subjected to meaningful one-to-one transformations so that the transformed space is still complete and each color is specified with a unique sum.

Elementary art education materials, dictionaries, and electronic search engines often define primary colors effectively as conceptual colors (generally red, yellow, and blue; or red, green, and blue) that can be used to mix "all" other colors and often go further and suggest that these conceptual colors correspond to specific hues and precise wavelengths. Such sources do not present a coherent, consistent definition of primary colors since real primaries cannot be complete.

Tetrachromacy

Tetrachromacy is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms with tetrachromacy are called tetrachromats.

In tetrachromatic organisms, the sensory color space is four-dimensional, meaning that to match the sensory effect of arbitrarily chosen spectra of light within their visible spectrum requires mixtures of at least four primary colors.

Tetrachromacy is demonstrated among several species of bird, fish, amphibian, reptile, insect and some mammals. It was the normal condition of most mammals in the past; a genetic change made the majority of species of this class eventually lose two of their four cones.

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 explanation of trichromacy is that the organism's retina contains three types of color receptors (called cone cells in vertebrates) 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 cells may contribute to color vision.

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 wavelengths from about 380 to 740 nanometers. In terms of frequency, this corresponds to a band in the vicinity of 430–770 THz.

The spectrum does not contain all the colors that the human eyes and brain can distinguish. Unsaturated colors such as pink, or purple variations like magenta, for example, are absent because they can only be made from a mix of multiple wavelengths. Colors containing only one wavelength are also called pure colors or spectral colors.

Visible wavelengths pass largely unattenuated through the Earth's atmosphere via the "optical window" region of the electromagnetic spectrum. An example of this phenomenon is when clean air scatters blue light more than red light, and so the midday sky appears blue. The optical window is also referred to as the "visible window" because it overlaps the human visible response spectrum. The near infrared (NIR) window lies just out of the human vision, as well as the medium wavelength infrared (MWIR) window, and the long wavelength or far infrared (LWIR or FIR) window, although other animals may experience them.

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