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The Evolution of Human Skin Color by Annie Prud’homme-Généreux Life Sciences Quest University, Canada

Part I Skin Cancer
“Stop it!” called Tatiana, playfully. Her boyfriend, Zach, was inspecting her skin very carefully. “Look,” he answered her, his voice taking on a more serious tone. “Today a woman walked into the clinic for her annual physical. Everything about her seemed ne. She leads a balanced lifestyle, she eats well, she exercises: she’s healthy! But as she was about to leave, I noticed a mole on her arm. It had many of the warning signs of skin cancer. So, I removed the mole. is woman now has to wait for the lab results to see if it was cancerous. If it is, maybe we caught it early enough to treat it, and maybe not. Either way, her life is changed. I just want to make sure you don’t have any suspicious moles, okay?” Tatiana relented and allowed Zach to examine her skin. She asked: “Do only white people get skin cancer?” “No, people of all skin tone can get skin cancer, but it does occur more frequently in Caucasians.”

Questions
1. What are the causes of skin cancer? 2. Why are Caucasians more at risk of skin cancer than other populations? 3. At what age does skin cancer typically occur? Is the incidence of skin cancer greater in youth or old age?

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Part II Skin Pigmentation and UV Light
Why are human populations di erently pigmented? What caused the evolution of an array of di erent skin colors?

Humans Were Initially Lightly Pigmented
About seven million years ago, humans and chimpanzees shared a common ancestor. Since that time, the two species have evolved independently from one another. It is generally assumed that chimpanzees changed less over that time period than humans—because they have remained in their original environment. Chimpanzees are therefore often used as a surrogate to make inferences about the physical and behavioral attributes of our common ancestors. e skin of chimps is light and covered with hair. From this observation, it has been inferred that our earliest ancestor was also probably light-skinned and covered with hair. Since humans and chimps diverged, humans left the protection of trees and adapted to a new environment (the open savannah). is change in habitat required several adaptations. Life on the savannah provided little shade and so little protection from the sun, and required a more active lifestyle (i.e., hunting as opposed to picking fruits). It is also hypothesized that the social interactions and strategizing required for successful hunting favored the development of a large brain, which consumed a lot of energy and generated heat. An increased number of sweat glands and loss of body hair evolved to dissipate heat. is created a new problem, as the light skin became exposed and vulnerable to the sun’s damaging ultraviolet (UV) radiation.

Melanin: Natural Sunscreen
UV light is harmful to living organisms because it causes changes (i.e., mutations) in the DNA sequence. Skin cells that produced a pigment called melanin were advantaged because melanin is a natural sunscreen; it absorbs the energy of UV light and shields cells from the radiation’s harmful e ects. Such cells were favored in evolution and now all human skin cells can produce this pigment. People vary in their skin tone due to di erences in the distribution, quantity, size, and type of melanin found in their skin cells. As you might suspect, people with dark skin tend to have larger and more numerous melanin-containing particles in their skin. is provides protection from the sun’s UV rays. Many genes are known to a ect the production of melanin and cause skin color variation in humans. While skin color is an inherited characteristic, the fact that many genes code for this trait explains why children do not always exactly match their parents’ skin tone. Tanning is the process of producing more melanin in the skin in response to ultraviolet exposure, and does not require a change in the genetic code (if a parent gets a tan, the o spring will not be more pigmented).

Distribution of UV Light across the Globe e following image (Figure 1) represents a map of the world on which the UV-light Index has been superimposed. e latitudes are shown on the left (latitude helps de ne a location on Earth, speci cally how far north or south of the equator a site is).

Questions
4. Does the amount of UV light reaching the Earth vary in a predictable manner? If so, describe the pattern you observe. 5. What latitude receives the greatest amount of UV light? e least? 6. Based on these data, where might you expect to nd the most lightly pigmented and most darkly pigmented people on the planet? Be as speci c as you can. 7. Provide a rationale to your answer above (i.e., why did you think that more darkly pigmented people would be found in those areas)?

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NATIONAL CENTER FOR CASE STUDY TEACHING IN SCIENCE Figure 1. Global UV Index Forecast.

Source: Figure obtained from the National Oceanic and Atmospheric Administration. Graph retrieved 18 October 2009 from http://www.cpc. ncep.noaa.gov/products/stratosphere/uv_index/gif_ les/uvi_world_f1.gif. is U.S. Government material is not subject to copyright protection within the United States.

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Part III Distribution of Skin Tones across the Globe
Let’s examine whether our predictions were correct. Figure 2 shows the relationship between latitude and the average skin re ectance of populations located throughout the world. Skin re ectance is a measure of pigmentation. e more a skin re ects light, the lighter it is in tone. Figure 2. Relationship of skin re ectance to latitude.

Source: Panel B of Figure 2 in Barsh (2003). Graph originally captioned as “Summary of 102 skin re ectance samples for males as a function of latitude, redrawn from Relethford (1997).” © 2003 Public Library of Science. This is an open-access article distributed under the terms of the Public Library of Science Open-Access License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Questions
8. Interpret this graph and the trend it describes. a. Is skin re ectance randomly distributed throughout the globe? If not, how would you describe the pattern? b. Restate your ndings in terms of skin color and UV light (instead of skin re ectance and latitude). c. How closely do these ndings match the predictions of your hypothesis (Question 6)? d. Some populations have skin colors that are darker or lighter than predicted based on their location (their data point falls somewhere outside of the line shown in Figure 2). What might explain the skin color of these exceptional populations? Propose a few hypotheses. 9. Hypothesize why di erent skin colors have evolved. Based on what you know, what factor is most likely to exert a selective pressure on skin color?

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Part IV Natural Selection and Evolution of Skin Color
Based on the information provided so far, it seems reasonable to hypothesize that darker skin evolved to protect against the harmful e ects of UV light. In particular, individuals who lacked optimal pigmentation for tropical latitudes had a greater risk of skin cancer and death. Until fairly recently, this was the leading hypothesis about the evolution of skin color. However, there is a problem with this hypothesis. Let’s see if you can nd it. Here is some basic information on evolution by natural selection. Evolution is a change in the gene pool of a population of organisms from generation to generation. Natural selection is but one of several mechanisms by which evolution can take place. rough natural selection, populations evolve and become adapted to their speci c environment. Natural selection will occur if the following three conditions are present: • Variation: e organisms in the population vary with regard to a trait. • Heredity: Variation in the trait has a genetic component transmissible to o spring. • Selective Pressure & Di erential Reproductive Success: Some traits increase the odds of surviving to reproductive age and successfully producing and rearing o spring in a given environment. Such traits are more adaptive. ose organisms having the better adapted trait leave more o spring behind—they are “naturally selected.” In the next generation, this adaptive (and inherited) trait will increase in frequency and will be represented in a greater proportion of the population. At this point, the genetic makeup of the population is di erent from that of the starting population: the population has evolved. Evolution is really a “number’s game”: the organisms that reproduce the most “win” because their traits will be disproportionally represented in the next generation. Note also that individuals do not evolve. ey either breed more e ectively or less e ectively, depending on already existing di erences in their traits. Only populations evolve or change over time.

Questions
10. Review your answer to Question 3. Keeping your answer in mind, how strong a selective pressure do you expect skin cancer (UV-induced mutations) to exert on reproductive success? 11. Based on this information, does your hypothesis about the evolution of skin color (Question 9) seem likely? Why or why not? How does skin color meet, or fail to meet, the three requirements of natural selection outlined above?

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Part V Folate: A Di erent Way of Looking at It
Since skin cancer tends to occur after age 50, it has little impact on reproductive success. Consequently, skin cancer probably exerted little pressure on the evolution of skin color. Some other factor must explain the range in pigmentation that is observed in the human population. For years, this fact was overlooked by the scienti c community, and the consensus was that dark skin had evolved as protection against skin cancer. In 1991, the anthropologist Nina Jablonski was skimming though scienti c journals when she came upon a 1978 paper by Branda and Eaton. is paper investigated the e ects of sunlight on an essential chemical found in our body: folate or folic acid (one of the B vitamins). Folate is an essential nutrient for DNA synthesis. Since cells reproduce at a fast pace during pregnancy (and hence, there is a lot of DNA replication), the highest levels of folate are needed during pregnancy. Folate de ciencies during pregnancy can lead to anemia in the mother and malformations of the nervous system (neural tube defects in particular), gastrointestinal system, aorta, kidney, and skeletal system in the fetus. ere is also a high rate of miscarriages. In addition, folate de ciency has been linked to spermatogenesis defects (inability to form sperm) in mice and rats (Mathur et al., 1977), and anti-folate agents are being investigated as a form of male contraceptive (Cosentino et al., 1990). Branda & Eaton’s paper measured the folate concentration in two human test groups. e results are shown in Figure 3. One group (called “patients”) was exposed to UV-light, while “normals” were not so exposed. Figure 3. Levels of blood folate in people exposed and not exposed to UV light.

Question
12. Based on Branda and Eaton’s results (Figure 3), what is the apparent e ect of UV light exposure on blood folate levels? Folate was isolated from blood and placed in a test tube. Half of the test tubes were exposed to UV light for 1 hour. e folate concentration in the samples was measured. e results are indicated in Table 1. Table 1. Folate concentrations in four samples of human plasma before and after a 1 hr exposure to UV light in vitro. Patients were exposed to UV light for at least 9 hours every day for 3 months. e di erence between the two groups was statistically signi cant (P< 0.005). Brackets represent the standard error of the mean.
Source: Figure 1 from Branda, R.F., and Eaton, J.W.(1978). Skin color and nutrient photolysis: An evolutionary hypothesis. Science 201: 625–626. Reprinted wih permission from AAAS. is gure and Table 1 may be used for non-commercial and classroom purposes only. Any other uses require the prior written permission from AAAS.

Source: Table 1 from Branda, R.F., and Eaton, J.W.(1978). Skin color and nutrient photolysis: An evolutionary hypothesis. Science 201: 625–626. Reprinted with permission from AAAS.

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Question
13. What is the apparent e ect of UV light on folate levels in these test tubes? Folate levels in humans are determined by two things: (1) dietary intake and (2) destruction through alcohol consumption or ultraviolet skin exposure.

Questions
How is folate linked to natural selection? All other things being equal, which skin tone would you expect to be correlated with higher levels of folate? Based on this new information, revise your hypothesis to explain the evolution of human skin color. What would happen to the reproductive success of: a. A light-skinned person living in the tropics? b. A light-skinned person living in the polar region? c. A dark-skinned person living in the tropics? d. A dark-skinned person living in the polar region? 18. Predict the skin tones expected at di erent latitudes, taking folate needs into consideration. Use the world map (Figure 4) to indicate the skin tone expected at each latitude (shade the areas where populations are darkly pigmented). Figure 4. Map of the world. 14. 15. 16. 17.

Source: http://commons.wikimedia.org/wiki/File:World_map_blank_black_lines_4500px_monochrome.png, CC BY-SA 3.0.

19. Can folate explain the variation and distribution of light- and dark-skinned individuals around the world?

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Part VI – Vitamin D: Still Another Way of Looking at It
Folate can explain why dark skin evolved, but it cannot account for the evolution of light skin. Another factor must be at play. Vitamin D3 is essential for normal growth, calcium absorption, and skeletal development. It is particularly important in maintaining and repairing healthy bones and teeth. Its role in calcium absorption makes it essential in maintaining a healthy heart, blood clotting, a stable nervous system, and an e ective immune system. De ciencies manifest themselves as rickets (softening of the bones), osteoporosis, and osteomalacia. It can lead to death, immobilization, or deformities. Women have a higher need for this nutrient during pregnancy and lactation due to their need to absorb calcium to build the fetal skeleton. Humans can obtain vitamin D3 by one of two means. ey can consume it in certain foods ( sh liver oil and to a lesser extent, egg yolk). Alternatively, skin cells have the ability to synthesize it from a cholesterol-like precursor. However, this process requires the energy of UV radiation. eoretical research on the dose of ultraviolet radiation required to produce vitamin D3 suggests that for moderately to darkly pigmented individuals (Figure 5): • ere is enough sunlight reaching the tropics (approximately 5° north of the Tropic of Cancer to approximately 5° south of the Tropic of Capricorn) to meet all of a human’s requirement for vitamin D3 throughout all months of the year. is is indicated by the dotted area on the map. Note: Vitamin D3 is not produced to toxic levels when high quantities of sunlight are present. • In the area indicated by narrowly-spaced obliques, there is not enough ultraviolet light to synthesize vitamin D3 in human skin for at least 1 month of the year; • In the area indicated by widely-spaced obliques, there is not enough UV light for the skin to synthesize vitamin D3 in any month of the year Figure 5. Amount of UV light available to synthesize recommended levels of vitamin D for a moderately to darkly pigmented person at various locations around the world.

Source: reprinted from e Journal of Human Evolution 39(1), Jablonski, N.G., and G. Chaplin, e Evolution of human skin coloration, pp. 57–106, Figure 2, copyright (2000), with permission from Elsevier. http://www.sciencedirect.com/science/journal/00472484. “ e Evolution of Human Skin Color” by Annie Prud’homme-Généreux Page

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Questions
20. How is vitamin D linked to natural selection? 21. Which skin tone allows someone to maintain the recommended level of vitamin D? 22. Based on this new information, revise your hypothesis to explain the evolution of the variation and distribution of human skin color. 23. Taking only vitamin D into consideration, what would happen to the reproductive success of: a. A light-skinned person living in the tropics? b. A light-skinned person living in the polar region? c. A dark-skinned person living in the tropics? d. A dark-skinned person living in the polar region? 24. Predict the skin tones expected at di erent latitudes, taking only vitamin D needs into consideration. Use the world map (Figure 6) to indicate the skin tone expected at each latitude (shade a region to represent pigmented skin in that population). Figure 6. Map of the world.

Source: http://commons.wikimedia.org/wiki/File:World_map_blank_black_lines_4500px_monochrome.png, CC BY-SA 3.0.

25. Can vitamin D alone explain the current world distribution of skin color? Evolution by natural selection is a process of compromise in which costs are minimized and bene ts are maximized. Both light and dark skins have costs and bene ts. As you are probably now realizing, adopting one level of pigmentation has trade-o s.

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NATIONAL CENTER FOR CASE STUDY TEACHING IN SCIENCE 26. Using principles of natural selection, predict the skin tone expected at di erent latitudes, taking ultraviolet exposure, vitamin D, and folate needs into consideration. Use the map (Figure 7) to indicate skin tone patterns at di erent latitudes (shade regions where populations are expected to be darkly pigmented). Figure 7. Map of the world.

Source: http://commons.wikimedia.org/wiki/File:World_map_blank_black_lines_4500px_monochrome.png, CC BY-SA 3.0.

27. Are UV light, vitamin D and folate needs su cient to explain the current world distribution of skin color? 28. How might you explain that Inuits, living at northern latitudes, are relatively dark-skinned (much more so than expected for their latitude)? Propose a hypothesis. 29. Conversely, Northern Europeans are slightly lighter-skinned than expected for their latitude. Propose a hypothesis to explain this observation.

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Part VII – Adaptation in Progress
While reading the newspaper, you stumble upon an article called “Are you getting enough vitamin D” (Mittelstaedt, 2007). It describes the results of a recent study on the vitamin D blood levels of students at the University of Toronto. e results are analyzed by the students’ ancestry. Intrigued by these ndings, you obtain and read the original research paper (Gozdzik et al., 2008). e gure below shows the levels of vitamin D found in the blood of Canadian students, shown by their ancestry (Gozdzik et al., 2008). e bottom of each box represents the 25th percentile of the data, the top of the box represents the 75th percentile, and the horizontal bar within the box is the median (the value below which 50% of all data points fall). 25-hydroxyvitamin D [25(OH)D] levels greater than 75 nmol/L are considered optimal (Holick & Chen, 2008). Note that similar results would be found in the United States (Calco & Whiting, 2003; Bodnar et al., 2007). Figure 8. Amount of vitamin D in blood of Canadians, shown by ancestry.

Source: Gozdzik et. al. 2008. © 2008 Gozdzik et al., an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Questions
30. Which populations have insu cient amounts of vitamin D in their blood? 31. What is a common trait to all of these populations? How could this explain their de ciency? 32. For several decades, milk and cereals have been forti ed with vitamin D to help increase intake of this nutrient. Hypothesize why these forti cation programs appear to be failing in some ethnic groups (there may be di erent reasons for di erent groups). 33. What should some Canadians do to avoid vitamin D de ciency?

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NATIONAL CENTER FOR CASE STUDY TEACHING IN SCIENCE 34. If individuals do not take steps to boost their vitamin D intake, hypothesize what might happen to their descendants in the future. 35. What might happen to Australians of European descent over time? 36. ink of the forces that a ected the evolution of skin color in the past. Contrast these to the factors that a ect skin pigmentation today. Hypothesize about the factors that may a ect skin pigmentation in the future. a. Do you think the impact of natural selection on skin color is as strong today as it was in the past? b. What factors may have decreased or increased selection today? c. Predict skin color distribution in the year 2500. Outline your assumptions and the reasons for your predictions. To beat the “winter-blahs,” a person of European descent is planning a trip to the Caribbean. Consider all the evidence you have learned in this case and apply it to this situation. What would you advise this lightly-pigmented person to do on the trip?

Questions
37. During the trip, the lightly-pigmented individual is looking forward to lying on a beach and working on his/ her tan. Will this person be adapting to the environment (in a Darwinian sense)? In other words, will evolution take place by developing a tan in a geographical region of intense UV light? 38. Should the lightly-pigmented person wear sunscreen on the trip? 39. Should this person wear sunscreen when he/she is at northern latitudes (i.e., at home in Canada)?

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Part VIII – Sexual Selection Too?
Sexual selection is a special type of natural selection. Darwin was the rst to recognize that if males or females of a species choose their mate based on a particular trait, then their preference will exert selective pressure on that trait. e result is that the opposite sex will evolve to meet that preference. An obvious example is the peacock’s tail. If females prefer males with the longest and most ornate and iridescent tail as mates, even if having such a tail seems detrimental to the male’s survival, it may be selected for evolution. is is because reproductive success, and not survival per se, is the most important characteristic that drives evolution. Females might prefer males with elaborate traits because it is a real signal of health and genetic worth: only males with good genes can a ord to have such a handicap (such a large tail). is process is called sexual selection. It occurs when individuals in the population di er in their ability to attract a mate. Research by Nina Jablonski and George Chaplin suggest that women generally produce 3–4% less melanin in their skin than do men in all populations of the world.

Questions
40. Could this observation be the result of sexual selection? Explain your answer. 41. Could this observation be the result of “normal” natural selection (i.e., not sexual selection)? Explain your answer.

References
Barsh, G.S. (2003). What controls variation in skin color? PLOS 1(1): 019–022. Bodnar, L.M., Simhan, H.N., Powers, R.W., Frank, M.P., Cooperstein, E., and Roberts, J.M. (2007). High prevalence of vitamin D insu ciency in black and white pregnant women residing in the northern United States and their neonates. Journal of Nutrition 137(2):447–52. Branda, R.F., and Eaton, J.W. (1978). Skin colour and nutrient photolysis: An evolutionary hypothesis. Science 201: 625–626. Calvo, M.S., and Whiting, S.J. (2003). Prevalence of vitamin D insu ciency in Canada and the United States: Importance to health status and e cacy of current food forti cation and dietary supplement use. Nutrition Reviews 61(3): 107–13. Cosentino, M.J., Pakyz, R.E., and Fried, J. (1990). Pyrimethamine: An approach to the development of a male contraceptive. Proceedings of the National Academy of Scences. (U.S.A.) 87, 1431–1435. Freeman, S. (2005). Biological Science, 2nd edition. Upper Saddle River, NJ: Pearson Prentice Hall. Gozdzik, A., Barta, J.L., Wu, H., Wagner, D., Cole, D.E., Vieth, R., Whiting, S., and Parra, E.J. (2008). Low wintertime vitamin D levels in a sample of healthy young adults of diverse ancestry living in the Toronto area: Associations with vitamin D intake and skin pigmentation. BMC Public Health 8: e336. Retrieved 22 April 2011 from http://www.biomedcentral.com/1471-2458/8/336#B37. Holick, M.F., and Chen, T.C. (2008). Vitamin D de ciency: A worldwide problem with health consequences. Am J Clin Nutr 2008, 87(4):1080S-1086S. Mathur, U., Datta, S.L., and Mathur, B.B. (1977). e e ect of aminopterin-induced folic acid de ciency on spermatogenesis. Fertility Sterility 28, 1356–1360. Mittelstaedt, M. (2007). Are you getting enough vitamin D. Globe and Mail Dec 19 2007. Retrieved 8 December 2009 from http://www.theglobeandmail.com/life/article804980.ece. Relethford, J.H. (1997). Hemispheric di erence in human skin color. Am J Phys Anthropol 104: 449–457.

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