Part Ia
... continued from Part I
It should also be noted that many humans carry more than one copy of the middle wavelength-sensitive cone opsin. As this is grist for the evolution of color vision mill, we're literally ripe for the addition of a fourth cone class. (This probably won't happen, though, because people with a fourth cone class will be constantly trying to readjust the color on television sets. As a result of that such people will be highly selected against in bars the world over
Since 1990, a few other opsins have been sequenced, specifically opsins from a variety of monkeys. I don't know as that they've been compared with the others, but I'm willing to predict where they should fit into the picture. It's nice to have a theory that let's you do that. (Since I wrote this last paragraph, I've seen another phylogeny that I think had more than twice as many opsin sequences as my best current reference. As far as I know, that work is still in press, and I no longer have access to it. From what I saw, though, the creationists have even more reason to fold their hand on this one now than they did two years ago...)
Comparative Psychology
Prior to the advent of some nifty techniques in molecular biology, people had to use less direct methods of classifying photoreceptors. Among these methods are: direct measurement of the absorptive properties of individual receptors, measurement of the electrical responses of cells to monochromatic lights, and the conditioning of learned behaviors. Thus even without molecular biology, we knew (and know) a lot about the pigments underlying color vision systems.
Based on this sort of information, it's clear that most vertebrates have at least two cone classes. In fact, many birds, turtles and fish have four or five. Many invertebrates are similarly well endowed, and last I heard, the mantis shrimp was the winner of the contest of who has the largest number of photoreceptor classes. Given that coral reef animals and tropical birds often appear very colorful to us, it's not surprising that they have well developed systems of color vision. That different animals have different numbers of receptor classes already tells us that color vision systems are not all equivalent (as Bob might have us believe). If we restrict ourselves to animals which have the same number of receptor classes, might we expect that their color vision systems are equivalent?
The answer is a resounding no. Let's compare the color vision systems of two animals that both have three photopic (e.g. active under bright illumination) photoreceptor classes. One is the human, the other is the honey bee (specifically the worker--I don't know how the other castes are endowed). Does anybody here think that what a bee sees when it looks at a rainbow has the same appearance as what we see? We'll ignore optical polarization (which the bee is sensitive to and we're not) and focus on what we can infer about "color" based on, among other things, our knowledge of the bee's receptor classes. To begin with, at the inside of the rainbow where the violet-appearing light fades off to invisibility for us, the bee will still see more rainbow. On the outside, where we see red, the bee would see nothing for although bees have an ability to see what for us is UV, we have the ability to see what bees might call infrared.
Now picture that rainbow: what you see appears to have discrete bands of color. Don't for a minute think that those bands arise from there being anything discrete about the radiation emanating from that patch of sky. If you measured the radiation with a spectrophotometer, you'd find that the wavelength of maximum intensity as a function of the radial distance across the rainbow would decrease smoothly and monotonically from the outside to the inside of the bow. The apparent discreteness is an artifact of our photopigments (chromophore + opsin) and the neural processing of our photoreceptors' outputs. The bee too would probably see discrete bands (We can't ever really know how the world appears to a bee, but given what we can infer from doable experiments -- I actually chose the bee in part because its color vision has been studied about as much as any other animal's, excluding the human's -- the supposition that it would see discrete "color" bands from a rainbow is reasonable.) However, just as the outer and inner borders would be in different locations for us and bees (as described in the preceding paragraph), the borders of each "color" would be placed differently by the bee as well.
I can't claim that we have a good handle on why different animals have different visual pigments. There are some cases that are well understood--most notably it was predicted some 20 years before verification that marine fish that live just above the aphotic zone would have only one pigment, and that that one pigment would have a maximal sensitivity down around 450 nm (for us light at this wavelength would appear blue). It makes sense that if there isn't much light around, an animal's photoreceptors will be adapted to respond most strongly to the wavelengths of light most readily available. Bioluminescent fish and insects also tend to have pigments that are adapted for maximal sensitivity to the wavelengths of light emitted by their photophores (the molecules responsible for the emission of light e.g. from the abdomens of fireflies). The specifics of what selective advantage other pigments in other environments might convey are still somewhat mysterious
One thing is clear, however. The best known predictor of what sort of pigments will be expressed by any given animal, is the pigments expressed by its nearest living relatives. To an evolutionary biologist this makes a lot of sense, of course.
There are a lot of other differences (or similarities) between manifestations of color vision systems in different animals. I've chosen to stick to a discussion of pigments here partly for simplicity, and partly because the straightforwardness of analyzing retinal receptors makes this the facet of color vision about which the most data is available. The point of this post is to say that it makes no sense to use the presence or absence of color vision in determining a phylogeny. If you want to be serious about asking what color vision and evolution have to say about each other, you have to ask specific questions about what sort of color vision different animals have.
go to Part II