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 tetrachromacy are called tetrachromats.

In tetrachromatic organisms, the sensory color space is four-dimensional, meaning that matching 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 birds, fishes, amphibians, and reptiles. The common ancestor of all vertebrates was a tetrachromat, but mammals evolved dichromacy, due to the nocturnal bottleneck, losing two of their four cones. Trichromats can see approximately 100 million colour combinations, but a tetrachromat can see more than a billion color combinations.

Physiology

The normal explanation of tetrachromacy is that the organism's retina contains four types of higher-intensity light receptors (called cone cells in vertebrates as opposed to rod cells, which are lower-intensity light receptors) with different spectral sensitivity. This means that the organism may see wavelengths beyond those of a typical human's vision, and may be able to distinguish between colors that, to a normal human, appear to be identical. Species with tetrachromatic color vision may have an unknown physiological advantage over rival species.

Humans

Apes (including humans) and Old World monkeys normally have three types of cone cell and are therefore trichromats. However, human tetrachromacy may be possible in some situations.

Tetrachromacy requires that there be 4 independent photoreceptor cell classes with different spectral sensitivity. However, there must also be the appropriate post-receptoral mechanism to compare the signals from the four classes of receptors. According to the opponent process theory, humans have three opponent channels, which grant trichromacy. Whether a fourth opponent channel is available to facilitate tetrachromacy is unclear.

Mice, which normally have only two cone pigments (and therefore two opponent channels), have been engineered to express a third cone pigment, and appear to demonstrate increased chromatic discrimination, possibly indicating trichromacy and suggesting they were able to create or re-enable a third opponent channel. This would support the theory that humans should be able to utilize a fourth opponent channel for tetrachromatic vision. However, the original publication's claims about plasticity in the optic nerve have also been disputed.

Tetrachromacy in carriers of CVD

It has been theorized that females who carry recessive opsin alleles that can cause color vision deficiency (CVD) could possess tetrachromacy. Female carriers of anomalous trichromacy (mild color blindness) possess heterozygous alleles of the genes that encode the L-opsin or M-opsin. These alleles often have a different spectral sensitivity, so if the carrier expresses both opsin alleles, they may exhibit tetrachromacy.

In humans, two cone cell pigment genes are present on the X chromosome: the classical type 2 opsin gene OPN1MW. People with two X chromosomes could possess multiple cone cell pigments, perhaps born as full tetrachromats who have four simultaneously functioning kinds of cone cell, each type with a specific pattern of responsiveness to different wavelengths of light in the range of the visible spectrum. One study suggested that 15% of the world's women might have the type of fourth cone whose sensitivity peak is between the standard red and green cones, giving, theoretically, a significant increase in color differentiation. Another study suggests that as many as 50% of women and 8% of men may have four photopigments and corresponding increased chromatic discrimination compared to trichromats. In 2010, after twenty years' study of women with four types of cones (non-functional tetrachromats), neuroscientist Gabriele Jordan identified a woman (subject ''cDa29'') who could detect a greater variety of colors than trichromats could, corresponding with a functional tetrachromat (or true tetrachromat).

Variation in cone pigment genes is wide-spread in most human populations, but the most prevalent and pronounced tetrachromacy would derive from female carriers of major red/green pigment anomalies, usually classed as forms of "color blindness" (protanomaly or deuteranomaly). The biological basis for this phenomenon is X-inactivation of heterozygotic alleles for retinal pigment genes, which is the same mechanism that gives the majority of female new-world monkeys trichromatic vision.

In humans, preliminary visual processing occurs in the neurons of the retina. It is not known how these nerves would respond to a new color channel, that is, whether they could handle it separately or just combine it in with an existing channel. Visual information leaves the eye by way of the optic nerve; it is not known whether the optic nerve has the spare capacity to handle a new color channel. A variety of final image processing takes place in the brain; it is not known how the various areas of the brain would respond if presented with a new color channel.

Tetrachromacy may also enhance vision in dim lighting, or in looking at a screen.

Conditional tetrachromacy

Despite being trichromats, humans can experience slight tetrachromacy at low light intensities, using their mesopic vision. In mesopic vision, both cone cells and rod cell