Color vision
Color vision (CV), 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 == Isaac Newton 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. The visible light spectrum 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 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 (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: == 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 (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 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 typical 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 (see below), 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 criticis
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