Color vision

Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different frequencies independently of light intensity.
Color perception is a part of the larger visual system and is mediated by a 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 ultimately leading to higher cognitive functions in the brain. 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 the evolution of color vision within different animal taxa. In primates, 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

discovered that white light 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.File:Eyesensitivity.svg|thumb|upright=1.25|Photopic relative brightness sensitivity of the human visual system as a function of wavelength The visible light spectrum ranges from about 380 to 740 nanometers. Spectral colors 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 or ultraviolet, 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 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 much higher.
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 color vision. In brighter light, such as daylight, vision is photopic: light is detected by cone cells 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 cells. 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 colors 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 colors, and metallic colors.
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 obtained 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, which is defined by the number of primaries required to represent the color vision. This is generally equal to the number of photopsins expressed: a correlation that holds for vertebrates but not invertebrates. The common vertebrate ancestor possessed four photopsins plus rhodopsin, 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:

Dimension	Characteristic	Occurrence
Achromacy	lack of any color perception	Most skates
Monochromacy	1D color vision	Some mammals, including Pinnipeds, Cetaceans and Xenarthra
Dichromacy	2D color vision	Most mammals and a quarter of color blind humans
Trichromacy	3D color vision	Most Old World monkeys and apes, including most humans; possibly monotremes and some marsupials
Tetrachromacy	4D color vision	Most birds, reptiles and fish, and rarely in humans
Pentachromacy and higher	5D+ color vision	Rare in vertebrates

Physiology of color perception

Perception of color begins with specialized retinal cells known as cone cells. 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 apoprotein covalently linked to a light-absorbing prosthetic group: 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, medium, and long 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 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 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;
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 theory. The trichromatic theory, or Young–Helmholtz theory, proposed in the 19th century by Thomas Young and Hermann von Helmholtz, posits three types of cones preferentially sensitive to blue, green, and red, respectively. Others have suggested that the trichromatic theory is not specifically a theory of color vision but a theory of receptors for all vision, including color but not specific or limited to it. Equally, it has been suggested that the relationship between the phenomenal opponency described by Ewald Hering and the physiological opponent processes are not straightforward, making of physiological opponency a mechanism that is relevant to the whole of vision, and not just to color vision alone. 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 shows that cyan, rather than green, is the complement of red, and that magenta, rather than red, is the complement of green. It therefore also shows that the reddish-green color supposed to be impossible by opponent process theory is actually 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 newer theory proposed by Edwin H. Land, the Retinex Theory, is based on a demonstration of color constancy, which shows that the color of any surface that is part of a complex natural scene is to a large degree independent of the wavelength composition of the light reflected from it. Also the after-image produced by looking at a given part of a complex scene is also independent of the wavelength 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" than "red" light is magenta, so is the after–image of the same surface when it reflects more "red" than "green" light. This would seem to rule out an explanation of color opponency based on retinal cone adaptation.
According to Land's Retinex theory, color in a natural scene depends upon the three sets of cone cells separately perceiving each surface's relative lightness in the scene and, together with the visual cortex, assigning color based on comparing the lightness values perceived by each set of cone cells.