Chameleon Color Changing: A DEEP Dive

CHAMELEON COLOR CHANGE:

As I’m sure you know, Chameleons are capable of color change. And, when I first was interested in how they did this, I looked it up and was provided with the simple answer “Chameleons can change color due to their special chromatophores”. This is something most people interested in chameleons know, but I still feel as though using the word chromatophore as an explanation leaves you just as much in the dark as you were before. So, what I want to do here today is explain on a deep scientific level how chromatophores work.


VISIBLE LIGHT:

However, to understand how chromatophores work, you need to understand two things: how light works, and how human skin works. We’ll start with how light works. Light is a type of electromagnetic radiation, which is to say a wave that carries energy through space. Electromagnetic radiation can have different frequencies and thus, different energies. When electromagnetic radiation has any wavelength between 400 and 700 nm (or 1.63 eV – 3.26 eV of energy), it is called visible light because it can be picked up by the human eye. To show you how this works, let’s say we have a brick which appears red to us. The brick itself doesn’t have a color, but a molecular structure. When the sun emits its white light (white light is a combination of all visible light), and that light hits the brick, all colors aside from red are absorbed by the brick due to the brick’s chemical structure. When the red is reflected back to your eye, you find the brick to be red.


HUMAN SKIN:

Now, let’s look at this light in a more complex biological system. I, myself am Caucasian so my skin appears a kind of peach. A cell’s color is determined by the proteins they produce. Some proteins when hit by light reflect certain colors, making the protein appear those colors. When a cell has a sufficient amount of a given protein, it appears to be that color. “Skin cells” are keratinocytes, and they contain keratin which doesn’t reflect light at all. Therefore, keratinocytes are clear. However, within your mesh of keratinocytes, you also have some melanocytes (melanin-producing cells). Melanin reflects brown, black or yellow light and so, from a glance, my skin looks peach because of the small but not negligible amount of melanin in my skin compared to keratin. There are also other things that dictate your skin color such as carotene and hemoglobin, but I’m not going to talk about that. If you want to look into that, it’s very interesting stuff.

Now, if you want to talk about the organization of human skin, we have a top layer of keratinocytes, and after a few layers of this, you’ll find the bottom layer of skin – called the stratum basale which has keratinocytes in addition to the melanocytes.



CHROMATOPHORES ARE NOT THE SAME AS MELANOCYTES:

Let’s talk about chameleon skin, starting from the outermost layer. Chameleons, like people have keratinocytes on the outermost layers. As you go deeper past the outer epidermis (epidermis just means “protective layer of skin”), you’ll find the upper dermis which contains the superficial iridophores (We’ll talk about these later). Then, in the deeper dermis are the deep iridophores and xanthophores, erythrophores and melanophores. You may notice that never once in that explanation did, I use the word chromatophore, and that’s because the word chromatophore just describes all color-producing cells in amphibians, fish, reptiles, crustaceans and cephalopods (things like squids). You are a human and so you do not have chromatophores but instead the melanocytes we discussed earlier. Birds also have melanocytes.

So, let’s talk about the difference between the two. Melanocytes are one kind of cell, while chromatophores are a whole class of cells. The aforementioned iridophores, xanthophores, erythrophores and melanophores are all chromatophores. There are also other chromatophores not found in chameleons called cyanophores and leucophores. Chromatophores are very closely associated with the vertebrate lineage. They probably evolved around 450 to 480 MYA in the jawed fish – the Gnathostomata. Cephalopods and crustaceans evolved chromatophores independently, so we’ll ignore them, but all vertebrates at one point or another had chromatophores. Both birds and mammals evolved to also have the melanocytes and then eventually lost their chromatophores.

The biggest difference between the melanocytes and the chromatophores are in how the color is actually produced. We already talked about how melanocytes produce the pigment melanin which is how they appear colored. Chromatophores will also produce some pigments, but they also sometimes contain guanine nanocrystals arranged in lattices. When the light hits these crystals, the dispersed light is shown to be blue or turquoise. These crystals can also be dispersed or aggregated, and depending on their level of concentration, when light hits the crystals, the skin will appear to be different colors.
 
IRIDOPHORES:

So, we mentioned earlier that the chameleons have on the outermost layer, keratinocytes. These are clear. But these are not chromatophores, and the chromatophores I want to start with are the superficial iridophores. Iridophores have those guanine crystals we talked about earlier. The crystals are stacked in layers and when the space between the layers is altered, color will reflect differently. When the layers are close, blue light is reflected and when the layers are far, green light is reflected. So, this allows chameleons to modulate between green and blue colorations.

Also, while chameleons do modulate areas of skin to be more blue or green, they also have some regions of skin with generally high iridophore crystal concentration and other regions of skin with generally low iridophore crystal concentration. This is why some Panther Chameleons (Furcifer pardalis) for instance will have blue and green stripes. But, in general, some chameleons might just be greener while others are bluer regardless of their color changing and so we can say that a blue chameleon like a Nosy Be just has more compacted iridophore crystals while a green chameleon like an Ambilobe has the opposite.

Now, if you’re asking why your blue Beta fish (Betta splendens) can’t turn as green as your chameleon, the answer is that while all Iridophore-containing animals certainly can alter their crystal concentration, very few can do it to the same degree as Chameleons.

Everything I’ve talked about up to this point is true about the superficial iridophores: the iridophores of the upper dermis. There are another kind of iridophores: the deeper iridophores of the deeper dermis. These guys do something completely different. The only reason they are called Iridophores is because they don’t use pigmentation and have guanine nanocrystals. Deep Iridophores actually have very little to do with the color of chameleon skin and so, we won’t get too deeply into them here. They function more so to deflect infrared light (780nm – 1mm). This is not red light, but a type of electromagnetic radiation outside of the visible light spectrum, which is potentially damaging, so the deep iridophores exist to properly reflect this light.


XANTHOPHORES AND ERYTHROPHORES:

Now, the iridophores are not the only kind of chromatophores in chameleons, because chameleons also have things called xanthophores. These are very different from iridophores because they don’t have the guanine crystals. Instead, these cells have vesicles (sacs of molecules in the cell) containing pigments. These vesicles will either contain pteridine and carotenoid pigments. Both of these pigments are going to absorb blue to violet wavelengths (400 – 500 nm) and reflect orange to yellow wavelengths (550 – 600 nm). This is going to produce for the cells, a nice orange-yellow color.

Now, chameleons also have erythrophores, which I will also discuss here because of how similar they are to the xanthophores. If xanthophores are the yellow-orange cells, erythrophores are the red-orange cells. They contain the same kind of pigments as the xanthophores, only they will be slightly different so that they reflect red to orange wavelengths (580 – 650 nm).

Now, a word about these pigments. Pteridine isn’t the name of a pigment, it is a class of pigments. The biggest difference between the carotenoid and pteridine pigment groups is where they come from. Chameleons are fully capable of producing pteridines, but they are not capable of producing cartenoids. Cartenoids actually come from their diet. See, cartenoids are found in plants, and so, when an insect eats a plant, it holds onto the cartenoids, and when the chameleon eats the insect, it takes the cartenoids, delivering them to the xanthophores and erythrophores.

So yes, if you’re chameleon is eating a diet devoid of cartenoids, it’s colors will change. This is just one of the reasons why gutloading your chameleon is so important. Cartenoids are a very interesting subject, and I don’t have time to get into them here, but when you’re looking for what foods to gutload with, I suggest you look at cartenoids. A diet high in cartenoids will allow your chameleon to have some beautiful colors.

Now, don’t think that a Parson’s chameleon (Calumma parsonii) will turn bright red when you feed it more carotenoids. Because that’s not how xanthophores and erythrophores work. Yes, a red or yellow chameleon probably has a large amount of xanthophores and erythrophores, but if your chameleon is green, that doesn’t mean it is carotenoid deficient or is missing its xanthophores and erythrophores. Maybe these cells are just less concentrated, or maybe their pigments are operating at lower levels, or they are working, but are tinting the chameleon instead of changing the chameleon’s color entirely. For example, if you look up a picture of a Jackson’s Chameleon (Trioceros jacksonii) you will see a green chameleon, but a green chameleon with a clear yellow tint. Iridophores should clearly make this chameleon within the green-blue range, yet here is a chameleon not only green, but with a yellow tint to it. Maybe a chameleon was going to be turquoise, but due to the presence of xanthophores and erythrophores, that turquoise was pushed towards the warm colors, producing a nice green.

Now, these cells do allow chameleons to change color, but in a different way. While iridophores are capable of producing changes in seconds, xanthophores and erythrophores will take minutes to hours. If the pigment is dispersed around the cell, the cell will be more yellow. If the pigment is concentrated, the cell will be redder or orange.

Now, the question is: what is the point of all this color changing. Due to the presence of iridophores, xanthophores and erythrophores, chameleons can be a range of colors from red to blue. Why would a chameleon want to be any of these colors? Well, most chameleons prefer to stay around the middle – green. The reason for this is that this is the best color for camouflage.

However, when a chameleon is trying to communicate something – say a female is trying to tell a male that it is pregnant, or a male is trying to tell a female he’s interested, or a male is trying to scare away another male – the last thing the chameleon wants to do is camouflage, and so a chameleon will try to be anything but green. A chameleon capable of being blue will fire up its iridophores to be blue. A chameleon capable of being red will fire up its erythrophores to be red.


MELANOPHORES:

The last kind of chromatophore I’ll talk about are the melanophores. Melanophores contain melanin (yes, that melanin), and melanin is able to make a cell appear darker or lighter. When melanin is concentrated, a cell will be dark and when it is dispersed, a cell will be light. This is because melanin reflects very little light, absorbing most of it. So, when the sun hits a melanophore, if the melanophore has been fired up, it will absorb most of the light from that sun ray.

Melanophores obviously work for temperature control. If a chameleon is cold, its melanophores will try to absorb more light, making the chameleon darker. If the chameleon is warm, its melanophores will relax and try to reflect light, making the chameleon lighter. This is also why your chameleon is very light when it sleeps. The melanophores (and in fact all chromatophores) are relaxed, so the chameleon appears pretty light-skinned.



HOW CHROMATOPHORES ARE ARRANGED:

Now, the question is: how are these chromatophores arranged with one another? Well, I can’t really tell you because I don’t know for certain. There haven’t been that many studies on chameleon chromatophores. And if you’re asking how I was able to write five pages on chromatophores if we don’t have information on them, the answer is that I really looked at the chromatophores of closely related animals and essentially, made a guess based on the information I had. Don’t worry though, because the truth is I’m not the only one who does this. For example, the reason many psychological experiments are conducted on mice is because mice are pretty closely related to humans (the Euarchontoglires only diverged in the Paleogene), so we can roughly equate the biology of the two. When I did research on chromatophores, I looked at mostly Iguanians (the Iguania diverged in the Cretaceous) and some other squamates like Geckos (the Bifurcata only diverged in the Jurassic). It isn’t ideal, but we can generally say that an iridophore in a gecko probably works similar to an iridophore in a chameleon.

The problem is that when it comes to talking about how chromatophores are organized with one another is a little sketchier. It is much more possible that a chameleon has a different chromatophore arrangement than a Gecko. However, I don’t want to leave you with nothing, so I did some research, and it turns out that we actually have a study on the chromatophore arrangement of the lizard Agama atra which like chameleons belongs to the clade Acrodonta (diverged in the Cretaceous).

What the research shows is that the skin had an outer layer of keratinocytes, followed by a thin basal lamina, followed by xanthophores, iridophores and then erythrophores. Beneath the erythrophores were the muscles.
 
Nice write up. You should add your bibliography so others can follow how you created this

And there’s probably some cool articles I would enjoy 😁
 
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