The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and UV radiation during human expansion
Open accessAbstract
This review examines putative, yet likely critical evolutionary pressures contributing to human skin pigmentation and subsequently, depigmentation phenotypes. To achieve this, it provides a synthesis of ideas that frame contemporary thinking, without limiting the narrative to pigmentation genes alone. It examines how geography and hence the quality and quantity of UV exposure, pigmentation genes, diet-related genes, vitamins, anti-oxidant nutrients, and cultural practices intersect and interact to facilitate the evolution of human skin color. The article has a strong focus on the vitamin D-folate evolutionary model, with updates on the latest biophysical research findings to support this paradigm. This model is examined within a broad canvas that takes human expansion out of Africa and genetic architecture into account. A thorough discourse on the biology of melanization is provided (includes relationship to BH4 and DNA damage repair), with the relevance of this to the UV sensitivity of folate and UV photosynthesis of vitamin D explained in detail, including the relevance of these vitamins to reproductive success. It explores whether we might be able to predict vitamin-related gene polymorphisms that pivot metabolism to the prevailing UVR exposome within the vitamin D-folate evolutionary hypothesis context. This is discussed in terms of a primary adaptive phenotype (pigmentation/depigmentation), a secondary adaptive phenotype (flexible metabolic phenotype based on vitamin-related gene polymorphism profile), and a tertiary adaptive strategy (dietary anti-oxidants to support the secondary adaptive phenotype). Finally, alternative evolutionary models for pigmentation are discussed, as are challenges to future research in this area.
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Hakim, O. A., Hart, K. H., McCabe, P., Berry, J., Robertson, F., Rhodes, L. E., Spyrou, N. M., Alfuraih, A. A., & Lanham-New, S. A. (2016) · The Journal of Steroid Biochemistry and Molecular Biology · doi:10.1016/j.jsbmb.2016.03.025 · Cited by 24 · see citing works
Jones, P. R., Lucock, M. D., Chaplin, G., Jablonski, N. G., Veysey, M. J., Scarlett, C. J., & Beckett, E. (2020) · Genes & Nutrition · doi:10.1186/s12263-020-00663-3 · Cited by 24 · see citing works
Self-citationInfluence of skin melanisation and ultraviolet radiation on biomarkers of systemic oxidative stress
Shih, B. B.-J., Farrar, M. D., Vail, A., Allan, D., Chao, M., Hu, C., Jones, G. D. D., Cooke, M. S., & Rhodes, L. E. (2020) · Free Radical Biology and Medicine · doi:10.1016/j.freeradbiomed.2020.07.034 · Cited by 23 · see citing works
Lucock, M. D., Jones, P. R., Veysey, M. J., Thota, R. N., Garg, M. L., Furst, J. E., Martin, C., Yates, Z. R., Scarlett, C. J., Jablonski, N. G., et al. (2021) · American Journal of Human Biology · doi:10.1002/ajhb.23667 · Cited by 22 · see citing works
Self-citationBeckett, E., Jones, P. R., Veysey, M. J., Duesing, K., Martin, C., Furst, J. E., Yates, Z. R., Jablonski, N. G., Chaplin, G., & Lucock, M. D. (2017) · American Journal of Human Biology · doi:10.1002/ajhb.23010 · Cited by 21 · see citing works
Self-citationLucock, M. D., Jones, P. R., Martin, C., Yates, Z. R., Veysey, M. J., Furst, J. E., & Beckett, E. (2018) · Nutrition Reviews · doi:10.1093/nutrit/nuy013 · Cited by 21 · see citing works
Self-citationHuman biological variation and the “normal”
Graves, J. L. (2021) · American Journal of Human Biology · doi:10.1002/ajhb.23658 · Cited by 21 · see citing works
Evolutionary Approach to Medicine
Berlim, M. T., & Abeche, A. M. (2001) · Southern Medical Journal · doi:10.1097/00007611-200101000-00005 · Cited by 20 · see citing works
Unpacking Human Evolution to Find the Genetic Determinants of Human Skin Pigmentation
Quillen, E. E., & Shriver, M. D. (2011) · Journal of Investigative Dermatology · doi:10.1038/skinbio.2011.3 · Cited by 20 · see citing works
Rearrangement and depletion of folate in human skin by ultraviolet radiation
Hasoun, L., Bailey, S., Outlaw, K. K., & Ayling, J. E. (2015) · British Journal of Dermatology · doi:10.1111/bjd.13885 · Cited by 18 · see citing works
Wolf, S. T., Stanhewicz, A. E., Jablonski, N. G., & Kenney, W. L. (2018) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00501.2018 · Cited by 18 · see citing works
Cabrera, S., Benavente, D., Alvo, M., de Pablo, P., & Ferro, C. J. (2014) · Journal of Photochemistry and Photobiology B Biology · doi:10.1016/j.jphotobiol.2014.07.001 · Cited by 17 · see citing works
The color of normal: How a Eurocentric focus erases pigmentation complexity
Norton, H. (2020) · American Journal of Human Biology · doi:10.1002/ajhb.23554 · Cited by 14 · see citing works
Darwinian Medicine's Drawn-Out Dawn
Pennisi, E. (2011) · Science · doi:10.1126/science.334.6062.1486 · Cited by 11 · see citing works
Vitamin-related phenotypic adaptation to exposomal factors: The folate-vitamin D-exposome triad
Lucock, M. D. (2021) · Molecular Aspects of Medicine · doi:10.1016/j.mam.2021.100944 · Cited by 10 · see citing works
Self-citationVitamin D: Genetics, Environment & Health
Martin, C. E. (2014) · Journal of Food & Nutritional Disorders · doi:10.4172/2324-9323.1000155 · Cited by 9 · see citing works
Converging Evolutionary, Environmental and Clinical Ideas on Folate Metabolism
Jones, P. R., Beckett, E., Yates, Z. R., Veysey, M. J., & Lucock, M. D. (2016) · Exploratory Research and Hypothesis in Medicine · doi:10.14218/erhm.2016.00003b · Cited by 9 · see citing works
Self-citationProgram of the 86th Annual Meeting of the American Association of Physical Anthropologists
García-Donas, J. G., Scholl, A. R., Dalton, A., Paine, R. R., & Kranioti, E. F. (2017) · American Journal of Physical Anthropology · doi:10.1002/ajpa.23210 · Cited by 7 · see citing works
Medicine in the 21st century: towards a Darwinian medical epistemology.
Román‐Franco, A. A. (2009) · PubMed · Cited by 5 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Bose, A., Sengupta, M., Maitra, S., Bandyopadhyay, A. R., & Schaschl, H. (2026) · Frontiers in Genetics · doi:10.3389/fgene.2026.1870791 · Cited by 2 · see citing works
- Authors
- Lucock, M. D.
- Affiliations
- University of Newcastle Australia (AU)
- Corresponding author
- Lucock, Mark D.
- ORCID
- Lucock, Mark D.
- Venue
- American Journal of Biological Anthropology
- Publisher
- Wiley
- Published
- 2022-06-25
- Volume
- 180
- Issue
- 2
- Pages
- 252-271
- ISSN
- 2692-7691 (ISSN-L)
- PMC
- PMC10083917
- OpenAlex
- W4283519774
- Semantic Scholar
- 956c104cf5dad95a30d6eaf272887728f04e6968
- Cited by
- 35
- References
- 115
- Citations by year
- 2026: 12 · 2025: 12 · 2024: 6 · 2023: 5
- FWCI
- 0.69
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- American Journal of Biological Anthropology (publishedVersion, cc-by-nc)
- PubMed Central (publishedVersion, cc-by-nc)
- PubMed (publishedVersion)
- Figshare (submittedVersion, cc-by-nc)
- NOVA (University of Newcastle Australia) (submittedVersion, cc-by-nc)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › melanin and skin pigmentation
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Race, Genetics, and Society
- Health Sciences › Medicine › Dermatology › Skin Protection and Aging
- Keywords
- human skin pigmentation, depigmentation, vitamin D, folic acid, ultraviolet radiation, melanization, folate biosynthesis, tetrahydrobiopterin, DNA repair, UV exposure, genetic polymorphism, dietary antioxidants, reproductive success
- MeSH terms
- Antioxidants, Folic Acid, Humans, Skin Pigmentation, Ultraviolet Rays, Vitamin A, Vitamin D, Vitamin K, Vitamins
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
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