Is it true that color vision is sex-linked for all species with binary sexes? Is there an evolutionary significance to the fact that color vision is X-linked in humans? E.g., only female humans can be tetrachromats.
Human females do not benefit much from an extra cone class, as color discrimination is hardly affected in most tetrachromats. In fact, tetrachromats often show increased error rates in their color discrimination. In dichromatic New World monkeys the heterozygous females do functionally benefit from their extra cone as they gain trichromatic vision.
Anomalous trichromats have all of their three cone types to perceive colors, but one type of cone perceives light slightly out of alignment. There are three different types of effect produced depending upon which cone type is affected.
The different anomalous conditions are protanomaly (reduced sensitivity to red light), deuteranomaly (reduced green sensitivity - the most common type) and tritanomaly (reduced blue sensitivity - extremely rare). People with deuteranomaly and protanomaly are collectively known as red-green colour blind and they generally have difficulty distinguishing between reds, greens, browns and oranges. They also commonly confuse different types of blue and purple hues. Red and green cones are indeed coded on the X-chromosome.
One source of variation is the very common Ser180Ala polymorphism that accounts for two spectrally different red pigments and that plays an important role in variation in normal color vision as well as defective color vision. The most common source of variation is the existence of several types of red/green pigment chimeras. The red and green-pigment genes are arranged in an array on the X-chromosome with one red-pigment gene followed by one or more green-pigment genes. Recombination events have given rise to red/green hybrid genes and deletion of the green-pigment genes. Only the first two genes in the tandem are expressed. The severity of red-green color vision defects is inversely proportional to the difference between the wavelengths of maximal absorption of the photopigments encoded by these two genes.
Females who are heterozygous for red and green pigment genes that encode three spectrally distinct photopigments have the potential for enhanced color vision, as they are effectively tetrachromats (Deeb, 2005, Neitz et al., 1991). However, sensitive color-contrast testing on 43 tetrachromats has revealed that most of these females have no deviating color-discrimination whatsoever. 8 subjects showed relatively small effects, while only one showed a clear increased sensitivity in a narrow range of frequencies. It is believed that the human visual system is not plastic enough to cope with the extra spectral input. In fact, in the group there was an overall increase in error rates on some color tests (pseudoisochromatic plates, and Nagel anomaloscope color matching) (Jordan & Mollon, 1993).
In New World Monkeys, however, the situation is different. Squirrel monkeys are basically a dichromatic species, but two-thirds of the females are heterozygous, and gain trichromatic vision by expressing two of three possible alleles coding for pigments in the middle- to long-wave range of the spectrum. X-chromosome inactivation serves to segregate the alternative allelic products in different subsets of cones. The visual system of the heterozygous female is apparently plastic enough to take advantage of the presence of three classes of cone, because heterozygous monkeys have enhanced color selectivity in the red-green range that are impossible for all males and for homozygous females. This advantage perhaps enables the heterozygote to judge better the ripeness of fruit, or to find fruit or conspecifics (Jordan & Mollon, 1993).
Note that the emergence of trichromacy in humans and some other primates was the result of the red/green gene duplication. Trichromacy in primates was evolutionary selected for likely because of the enhanced capability to discern (ripe) fruits (Lucas et al., 2003). It has nothing to do with sex-differences, because not many human females benefit from tetrachromacy in terms of enhanced color vision.
- Deeb, Clin Genet (2005); 67: 369–377
- Jordan & Mollon, Vis Res (1993); 33(11): 1495-1508
- Lucas et al., Evolution (2003); 57(11): 2636–43
- Neitz et al., Science (1991); 252(5008): 971-4
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