Rainbow of Colors: Monkey See
Color vision is something I take for granted. I forget that a significant portion of the population does not see the range of colors that I can. And I have no way of appreciating the range of colors that birds and bees can see. The natural history of primate color vision makes for an interesting story.
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We see color because the cones in our retinas make three different versions of proteins called opsins. These proteins are responsible for initiating impulses to the brain, and they have differing sensitivities to different wavelengths of light. It is by comparing the signal strength between the three opsins that our brains are able to interpret color. Since we have three types of opsin proteins (S, M and L), we are trichromats.
(Figure shows spectral sensitivities of the opsins in different organisims [1].)
Although there are several types and causes of color blindness, the most familiar cause is genetic and mainly affects men. This is because two of the three opsin genes are located on the X chromosome. Since females have two X chromosomes it doesn't matter if one of them has a defective opsin gene. But if a male has a defective opsin on the X chromosome, he's out of luck.
Color vision appears to have been the norm for most animals--including fish--but most placental mammals are dichromats, apparently because they lost two unnecessary opsins while adopting a nocturnal lifestyle early in their evolution. Most primates are different, however, and this is where the story gets interesting.
There is a basic divide among the monkeys and apes: there are the Old World Monkeys (OWMs) and Apes (including humans), and then there are the New World Monkeys (NWMs). NWM males are dichromats, but some females are trichromats. How can this be? Well instead of having two different opsins on the X chromosome, there are different variants (allels) of the one opsin gene on the X chromosome circulating in the population. Since females have two X chromosomes, they can have two variants of that opsin at once. Such females are called 'allelic trichromats'. Males are stuck with either one or the other variant, and thus remain dichromats. The exception is the howler monkey, which at some point had a gene duplication occur such that the X chromosome has two copies of the opsin gene. One of the opsins mutated such that it has a slightly different wavelength sensitivity, making howler monkeys 'true trichromats,' like us.
OWMs and Apes are true trichromats. The reason for this is similar to the howler monkey. At an early point after the split from NWMs, a gene duplication occured on the X chromosome leading to the existence of two opsin genes of slightly different peak sensitivity. However this was an earlier and separate event from that of the howler monkey. (In the figure above and to the left [2], the arrows indicate lineages that are true trichromats. Red lineages indicate those with a high precentage of OR pseudogenes, as explained below. Click to enlarge.)
All of this is interesting for a number of reasons, including issues of phylogeny, biogeography and so forth, but the story goes one step further. A group of scientists were studying the olfactory receptors (ORs) in primates. These proteins are to smell, what opsins are to vision--except that there are a greater variety of ORs. Specifically, they looked at the proportion of OR genes that are broken (pseudogenes) in a variety of primates and found something interesting. OWMs and Apes have a higher percentage of OR pseudogenes than NWMs, with one exception: the howler monkey. The figure to the right (click to enlarge) shows the percentage of OR pseudogenes in different primate species [2]. Humans have the highest percentage, but note that the only species above 20% are true trichromats. It is too early to say whether there is a causal relationship, but the correlation is striking and suggests that as primates gained better color vision, selective pressure for smell was relaxed, which allowed increasing numbers of ORs to become broken and unused.
As I look out my window, fall colors are on full display. Limited as they may be in comparision to some other animals, I'm grateful for my senses.
1. Vorobyev, Ecology and Evolution of Primate Colour Vision (pdf), Clinical and Experimental Optometry, 2004, 87:4-5. The same issue of this journal has several additional articles on color vision, freely available here.
2. Gilad et al., Loss of Olfactory Receptor Genes Coincides with the Acquisition of Full Trichromatic Vision in Primates, PLoS Biology, 2004, Vol. 2 Issue 1.
See "The Howler Monkey's Tale" in Richard Dawkins' Ancestor's Tale for a popular-science treatment of primate color vision.