Opponent process
The opponent process is a hypothesis of color vision that states that the human visual system interprets information about color by processing signals from the three types of photoreceptor cells in an antagonistic manner. The three types of cones are called L, M, and S. The names stand for "Long wavelength sensitive,” "middle wavelength sensitive," and "short wavelength sensitive." The opponent-process theory implicates three opponent channels: L versus M, S versus (L+M), and a luminance channel (+ versus -). These cone-opponent mechanisms were at one time thought to be the neural substrate for a psychological theory called Hering's Opponent Colors Theory, which calls for three psychologically important opponent color processes: red versus green, blue versus yellow, and black versus white (luminance). The Opponent Colors Theory is named for the German physiologist Ewald Hering who proposed the idea in the late 19th century. However, it has been argued that Hering’s Opponent Colors Theory lacks adequate phenomenological and empirical support, and may not be a necessary feature of normal human color experience. Correspondingly, considerable physiological and behavioral evidence proves that the physiological cone opponent mechanisms do not constitute the neurobiological basis for Hering's Opponent Colors Theory. == Color theory == === Complementary colors === When staring at a bright color for a while (e.g. red), then looking away at a white field, an afterimage is perceived, such that the original color will evoke its complementary color (cyan, in the case of red input). When complementary colors are combined or mixed, they "cancel each other out" and become neutral (white or gray). That is, complementary colors are never perceived as a mixture; there is no "greenish red" or "yellowish blue", despite claims to the contrary. The strongest color contrast that a color can have is its complementary color. Complementary colors may also be called "opposite colors" and they were originally considered the primary evidence in support of Hering's Opponent Colors Theory. There are two fatal problems with this evidence. First, the complement of red is not green, as called for by Hering's theory; it is bluish-green. And second, there exists a complementary color for every color, so there is nothing special about the set of complementary pairs picked out by Hering's theory. === Unique hues === The colors that define the extremes for each opponent channel are called unique hues, as opposed to composite (mixed) hues. Ewald Hering first defined the unique hues as red, green, blue, and yellow, and based them on the concept that these colors could not be simultaneously perceived. For example, a color cannot appear both red and green. These definitions have been experimentally refined and are represented today by average hue angles of 353° (carmine red), 128° (cobalt green), 228° (cobalt blue), 58° (yellow). The unique hues are a defining feature of many psychological color spaces, but there is substantial evidence showing that the unique hues are not hard wired in the nervous system, contrary to the stipulations of Hering's Opponent Colors Theory. Unique hues can differ between individuals and are often used in psychophysical research to measure variations in color perception due to color-vision deficiencies or color adaptation. While there is considerable inter-subject variability when defining unique hues experimentally, an individual's unique hues are very consistent, to within a few nanometers of wavelength. == Physiological basis == === Relation to LMS color space === The trichromatic theory is in conflict with Hering's Opponent Colors Theory, although it is compatible with a physiological opponent process that compares the outputs of the different classes of cone types. The poles of these cone opponent mechanisms do not correspond to the unique hues of Hering's Opponent Colors Theory and unlike the unique hues, have no privilege in color perception. Most humans have three different cone cells in their retinas that facilitate trichromatic color vision. Colors are determined by the proportional excitation of these three cone types, i.e. their quantum catch. The levels of excitation of each cone type are the parameters that define LMS color space. To calculate the opponent process tristimulus values from the LMS color space, the cone excitations must be compared: The luminous (achromatic) opponent channel is a weighted sum of all three cone cells (plus the rod cells in some conditions). The red–green opponent channel is equal to the difference of the L- and M-cones. The blue–yellow opponent channel is equal to the difference of the S-cone and the average/weighted sum of the L- and M-cones. Most mammals have no L cone (the primate L cone arose from a gene duplication of the M cone opsin gene). These mammals still show two kinds of opponent channels in their retinal ganglion cells: the achromatic channel and the blue-yellow opponency channel. === Cone opponent mechanisms are encoded in the retina === The output of different types of cones are compared by cells in the retina including retina bipolar cells (which compare signals from L and M cones) and bistratified retinal ganglion cells (which compare S cone signals with L and M cone signals). The output of bipolar cells is relayed to the visual cortex by the retinal ganglion cells (RGCs) by way of a thalamic relay station called the lateral geniculate nucleus (LGN) of the thalamus. Much of the scientific knowledge of retinal ganglion cell physiology was obtained by neural recordings of cells in the LGN. The cone-opponent mechanisms in the retina and LGN represent a fundamental physiological opponent process but do not represent the unique hues (or Hering's Opponent Colors Theory). For example, the colors that best elicit responses of the bistratified S-(L+M)-opponent neurons are best described as purplish (or lavender) and lime-green, not "blue" and "yellow". The neurons are sometimes referred to as "blue–yellow" neurons, but this is a historical artifact dating to the time when it was thought that Hering's Opponent Colors Theory was hardwired by the retina and the mismatch between the colors to which they are optimally tuned and Hering's Opponent Colors was overlooked. Cone opponent mechanisms exist in the retinas of many mammals, including monkeys, mice, and cats. In primates, the LGN contains three major classes of layers: Magnocellular layers (M, large-cell) – responsible largely for the luminance channel Parvocellular layers (P, small-cell) – responsible largely for red–green opponency Koniocellular layers (K) – responsible largely for blue–yellow opponency, poor spatial resolution, long latency Other mammals such as cats also have three cell types denoted as X (magno), Y (parvo), and W (konio). The W type is beyond most doubt homologous to the primate K type. There are some subtle differences between the M and X types as well as the Y and P types to make the correspondence unclear. === Advantage === Transmitting information in opponent-channel color space could be advantageous over transmitting it in LMS color space ("raw" signals from each cone type). There is some overlap in the wavelengths of light to which the three types of cones (L for long-wave, M for medium-wave, and S for short-wave light) respond, so it is more efficient for the visual system (from a perspective of dynamic range) to record differences between the responses of cones, rather than each type of cone's individual response. Hurvich and Jameson argued that the use of opponent-channel color space would increase color contrast, making the information easier to process by later stages of vision. === Color blindness === Color blindness can be classified by the cone cell that is affected (protan, deutan, tritan) or by the opponent channel that is affected (red–green or blue–yellow). In either case, the channel can either be inactive (in the case of dichromacy) or have a lower dynamic range (in the case of anomalous trichromacy). For example, individuals with deuteranopia see little difference between the red and green unique hues. == History == Johann Wolfgang von Goethe first studied the physiological effect of opposed colors in his Theory of Colours in 1810. Goethe arranged his color wheel symmetrically "for the colours diametrically opposed to each other in this diagram are those which reciprocally evoke each other in the eye. Thus, yellow demands purple; orange, blue; red, green; and vice versa: Thus again all intermediate gradations reciprocally evoke each other." Ewald Hering proposed opponent color theory in 1892. He thought that the colors red, yellow, green, and blue are special in that any other color can be described as a mix of them, and that they exist in opposite pairs. That is, either red or green is perceived and never greenish-red: Even though yellow is a mixture of red and green in the RGB color theory, humans
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