Eye color (English Wikipedia)

Debrowski, Adam. "Which Eye Colors Are the Rarest?". All About Vision. Retrieved 4 February 2021.
"Amber eyes". All About Vision. Retrieved 9 May 2021.
"Gray eyes". All About Vision. Retrieved 9 May 2021.

antropologia-fizyczna.pl

"Martin-Schulz Eye Color Chart". Archived from the original on 2 August 2016. Retrieved 13 January 2017.

archive.org

Cavalli-Sforza, Luigi Luca; Cavalli-Sforza, Luca; Menozzi, Paolo; Piazza, Alberto (1994). The History and Geography of Human Genes. Princeton University Press. ISBN 978-0-691-08750-4.^{[page needed]}

athro.com

Morris, PJ. "Phenotypes and Genotypes for human eye colors." Athro Limited website. Retrieved 10 May 2006.

biology-online.org

Jones, S.L.; Schnirel, B.L. (2006). "Subspecies comparison of the Genus: Corucia". Polyphemos. 4 (1): 1–25. Archived from the original on 2 February 2009.

books.google.com

Fox, Denis Llewellyn (1979). Biochromy: Natural Coloration of Living Things. University of California Press. ISBN 978-0-520-03699-4.
Pilli, E.; Berti, A. (2021). Forensic DNA Analysis: Technological Development and Innovative Applications. Apple Academic Press. p. 207. ISBN 978-1-000-06454-4. Retrieved 22 May 2024. "Several research groups have demonstrated that females have a darker eye color than males, given the same SNP profile (Martinez-Cadenas et al., 2013; Pietroni et al., 2014; Pospiech et al., 2016)."
Fox, Denis Llewellyn (1979). Biochromy: Natural Coloration of Living Things. University of California Press. p. 9. ISBN 978-0-520-03699-4.
Beddoe, John (1971). The Races of Britain. A contribution to the anthropology of Western Europe. Michigan State University. p. 39. ISBN 9780091013707. the Saxons...being very often hazel-eyed
(in French) Provincia: bulletin trimestriel de la Société de Statistique ..., Volumes 16–17 By Société de statistique, d'histoire et d'archéologie de Marseille et de Provence p. 273 l'iris gris est celui des chaouias...

booksc.eu

ur.booksc.eu

Mitchell R, Rochtchina E, Lee A, Wang JJ, Mitchell P (2003). "Iris color and intraocular pressure: the Blue Mountains Eye Study". Am. J. Ophthalmol. 135 (3): 384–6. doi:10.1016/S0002-9394(02)01967-0. PMID 12614760.

boston.com

Belkin, Douglas (17 October 2006). "Don't it make my blue eyes brown Americans are seeing a dramatic color change". The Boston Globe. Archived from the original on 22 October 2014.

carnby.altervista.org

"Distribution of Bodily Characters. Pigmentation, the Pilous System, and Morphology of the Soft Parts". Archived from the original on 26 July 2011.

coalitionbrewing.com

"Where are hazel eyes most common?". 26 February 2023.

doi.org

Wielgus AR, Sarna T (2005). "Melanin in human irides of different color and age of donors". Pigment Cell Res. 18 (6): 454–64. doi:10.1111/j.1600-0749.2005.00268.x. ISSN 0893-5785. PMID 16280011.
Prota G, Hu DN, Vincensi MR, McCormick SA, Napolitano A (1998). "Characterization of melanins in human irides and cultured uveal melanocytes from eyes of different colors". Exp. Eye Res. 67 (3): 293–9. doi:10.1006/exer.1998.0518. PMID 9778410.
Huiqiong Wang; Lin, S.; Xiaopei Liu; Sing Bing Kang (2005). "Separating reflections in human iris images for illumination estimation". Tenth IEEE International Conference on Computer Vision (ICCV'05) Volume 1. pp. 1691–1698 Vol. 2. CiteSeerX 10.1.1.87.418. doi:10.1109/ICCV.2005.215. ISBN 978-0-7695-2334-7. S2CID 2215768.
Mason, Clyde W. (1924). "Blue Eyes". Journal of Physical Chemistry. 28 (5): 498–501. doi:10.1021/j150239a007.
Oliphant LW (1987). "Pteridines and purines as major pigments of the avian iris". Pigment Cell Res. 1 (2): 129–31. doi:10.1111/j.1600-0749.1987.tb00401.x. PMID 3507666.
Kayser, Manfred; Liu, Fan; Janssens, A. Cecile J.W.; Rivadeneira, Fernando; Lao, Oscar; Van Duijn, Kate; Vermeulen, Mark; Arp, Pascal; et al. (2008). "Three genome-wide association studies and a linkage analysis identify HERC2 as a human iris color gene". Am. J. Hum. Genet. 82 (2): 411–23. doi:10.1016/j.ajhg.2007.10.003. PMC 2427174. PMID 18252221.
White, Désirée; Rabago-Smith, Montserrat (14 October 2010). "Genotype–phenotype associations and human eye color". Journal of Human Genetics. 56 (1): 5–7. doi:10.1038/jhg.2010.126. PMID 20944644.
Duffy, David L.; Montgomery, Grant W.; Chen, Wei; Zhao, Zhen Zhen; Le, Lien; James, Michael R.; Hayward, Nicholas K.; Martin, Nicholas G.; Sturm, Richard A. (2007). "A three-single-nucleotide polymorphism haplotype in intron 1 of OCA2 explains most human eye-color variation". Am. J. Hum. Genet. 80 (2): 241–52. doi:10.1086/510885. PMC 1785344. PMID 17236130.
Sturm RA, Frudakis TN (2004). "Eye colour: portals into pigmentation genes and ancestry" (PDF). Trends Genet. 20 (8): 327–32. doi:10.1016/j.tig.2004.06.010. PMID 15262401. Archived from the original (PDF) on 9 September 2006.
Grant MD, Lauderdale DS (2002). "Cohort effects in a genetically determined trait: eye colour among US whites". Ann. Hum. Biol. 29 (6): 657–66. doi:10.1080/03014460210157394. PMID 12573082. S2CID 25364754.
Liu, Fan; Van Duijn, Kate; Vingerling, Johannes R.; Hofman, Albert; Uitterlinden, André G.; Janssens, A. Cecile J.W.; Kayser, Manfred (2009). "Eye color and the prediction of complex phenotypes from genotypes". Current Biology. 19 (5): R192 – R193. Bibcode:2009CBio...19.R192L. doi:10.1016/j.cub.2009.01.027. PMID 19278628.
Martinez-Cadenas, Conrado; Peña-Chilet, Maria; Ibarrola-Villava, Maider; Ribas, Gloria (2013). "Gender is a major factor explaining discrepancies in eye colour prediction based on HERC2/OCA2 genotype and the IrisPlex model". Forensic Science International: Genetics. 7 (4). Elsevier BV: 453–460. doi:10.1016/j.fsigen.2013.03.007. hdl:10234/86709. ISSN 1872-4973. PMID 23601698. "This effect is what may explain the fact that there seem to be comparatively higher frequencies of blue-eyed males than blue-eyed females in populations of European origin such as Iceland [10], Holland [10], Australia [18] or Poland [17], as well as in this study (see Fig. 2)."
Hernando, Barbara; Ibarrola-Villava, Maider; Fernandez, Lara P.; Peña-Chilet, Maria; Llorca-Cardeñosa, Marta; Oltra, Sara S.; Alonso, Santos; Boyano, Maria Dolores; Martinez-Cadenas, Conrado; Ribas, Gloria (2016). "Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin". Biology of Sex Differences. 7 (1): 17. doi:10.1186/s13293-016-0070-1. hdl:10810/32358. ISSN 2042-6410. PMC 4797181. PMID 26998216. The results of this study suggest that there are indeed sex-specific genetic effects in human pigmentation, with larger effects for darker pigmentation in females compared to males. A plausible cause might be the differentially expressed melanogenic genes in females due to higher oestrogen levels. These sex-specific genetic effects would help explain the presence of darker eye and skin pigmentation in females, as well as the well-known higher melanoma risk displayed by males.
Hanel, Andrea; Carlberg, Carsten (September 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 864–875. doi:10.1111/exd.14142. ISSN 0906-6705. PMID 32621306. S2CID 220335539. "The genetic history of today's European populations is based on continuous migrations over the past 40 000 years. Homo sapiens arrived in Europe from Near East some 42 000 years ago.[48, 49] Like in their African origin, these humans had dark skin but due to variations of their OCA2 gene (causing iris depigmentation) many of them had blue eyes[48,50] (Figure 2, left)." Figure 2: "Phenotype information was retrieved from supplementary files of the according publications or additionally assessed following the instructions of HirisPlex-S, which is a forensic DNA phenotyping tool based on an array of different marker SNPs providing additional support to the SNPs of the genes SLC24A5, SLC45A2 and OCA2."
Sulem, Patrick; Gudbjartsson, Daniel F; Stacey, Simon N; Helgason, Agnar; Rafnar, Thorunn; Magnusson, Kristinn P; Manolescu, Andrei; Karason, Ari; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nat. Genet. 39 (12): 1443–52. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549.
Liu, Fan; Wollstein, Andreas; Hysi, Pirro G.; Ankra-Badu, Georgina A.; Spector, Timothy D.; Park, Daniel; Zhu, Gu; Larsson, Mats; Duffy, David L.; Montgomery, Grant W.; MacKey, David A.; Walsh, Susan; Lao, Oscar; Hofman, Albert; Rivadeneira, Fernando; Vingerling, Johannes R.; Uitterlinden, André G.; Martin, Nicholas G.; Hammond, Christopher J.; Kayser, Manfred (2010). "Digital Quantification of Human Eye Color Highlights Genetic Association of Three New Loci". PLOS Genetics. 6 (5): e1000934. doi:10.1371/journal.pgen.1000934. PMC 2865509. PMID 20463881.
Bito, LZ (1997). "Eye Color Changes Past Early Childhood". Archives of Ophthalmology. 115 (5): 659–63. doi:10.1001/archopht.1997.01100150661017. PMID 9152135.
Oliphant LW (1987). "Observations on the pigmentation of the pigeon iris". Pigment Cell Res. 1 (3): 202–8. doi:10.1111/j.1600-0749.1987.tb00414.x. PMID 3508278.
Oliphant LW (1981). "Crystalline pteridines in the stromal pigment cells of the iris of the great horned owl". Cell Tissue Res. 217 (2): 387–95. doi:10.1007/BF00233588. PMID 7237534. S2CID 8061493.
Lefohn A, Budge B, Shirley P, Caruso R, Reinhard E (2003). "An Ocularist's Approach to Human Iris Synthesis". IEEE Comput. Graph. Appl. 23 (6): 70–5. doi:10.1109/MCG.2003.1242384. S2CID 537404.
Ruiz-Linares, Andrés; Adhikari, Kaustubh; Acuña-Alonzo, Victor; Quinto-Sanchez, Mirsha; Jaramillo, Claudia; Arias, William; Fuentes, Macarena; Pizarro, María; Everardo, Paola; de Avila, Francisco; Gómez-Valdés, Jorge; León-Mimila, Paola; Hunemeier, Tábita; Ramallo, Virginia; Silva de Cerqueira, Caio C. (25 September 2014). "Admixture in Latin America: Geographic Structure, Phenotypic Diversity and Self-Perception of Ancestry Based on 7,342 Individuals". PLOS Genetics. 10 (9): e1004572. Bibcode:2014PLOSG..10.4572R. doi:10.1371/journal.pgen.1004572. ISSN 1553-7390. PMC 4177621. PMID 25254375.
Zhu, Gu; Evans, David M.; Duffy, David L.; Montgomery, Grant W.; Medland, Sarah E.; Gillespie, Nathan A.; Ewen, Kelly R.; Jewell, Mary; Liew, Yew Wah; Hayward, Nicholas K.; Sturma, Richard A.; Trenta, Jeffrey M.; Martina, Nicholas G. (2004). "A genome scan for eye color in 502 twin families: most variation is due to a QTL on chromosome 15q". Twin Res. 7 (2): 197–210. doi:10.1375/136905204323016186. PMID 15169604.
Mitchell R, Rochtchina E, Lee A, Wang JJ, Mitchell P (2003). "Iris color and intraocular pressure: the Blue Mountains Eye Study". Am. J. Ophthalmol. 135 (3): 384–6. doi:10.1016/S0002-9394(02)01967-0. PMID 12614760.
Lindsey JD, Jones HL, Hewitt EG, Angert M, Weinreb RN (2001). "Induction of tyrosinase gene transcription in human iris organ cultures exposed to latanoprost". Arch. Ophthalmol. 119 (6): 853–60. doi:10.1001/archopht.119.6.853. PMID 11405836.
Regan S, Judge HE, Gragoudas ES, Egan KM (1999). "Iris color as a prognostic factor in ocular melanoma". Arch. Ophthalmol. 117 (6): 811–4. doi:10.1001/archopht.117.6.811. PMID 10369595.
Hawkins TA, Stewart WC, McMillan TA, Gwynn DR (1994). "Analysis of diode, argon, and Nd: YAG peripheral iridectomy in cadaver eyes". Doc Ophthalmol. 87 (4): 367–76. doi:10.1007/BF01203345. PMID 7851220. S2CID 30893783.
Hammond BR, Fuld K, Snodderly DM (1996). "Iris color and macular pigment optical density". Exp. Eye Res. 62 (3): 293–7. doi:10.1006/exer.1996.0035. PMID 8690039.
Muiños Díaz, Yerena; Saornil, Maria A.; Almaraz, Ana; Muñoz-Moreno, Mari F.; García, Ciro; Sanz, Ruperto (2009). "Iris color: validation of a new classification and distribution in a Spanish population-based sample". European Journal of Ophthalmology. 19 (4): 686–689. doi:10.1177/112067210901900427. ISSN 1120-6721. PMID 19551689. S2CID 40940828.
Keyser, Christine; Bouakaze, Caroline; Crubézy, Eric; Nikolaev, Valery G.; Montagnon, Daniel; Reis, Tatiana; Ludes, Bertrand (2009). "Ancient DNA Provides New Insights into the History of South Siberian Kurgan People". Human Genetics. 126 (3): 395–410. doi:10.1007/s00439-009-0683-0. PMID 19449030. S2CID 21347353. Indeed, among the SNPs tested was rs12913832, a single DNA variation within a regulatory element of HERC2 gene which is associated to blue eye color in humans. This polymorphism, together with the diplotypes obtained from variations of the OCA2 locus (major contributor to the human eye color variation) showed that at least 60% of the ancient Siberian specimens under study had blue (or green) eyes.
Sulem, P.; Gudbjartsson, D.; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nature Genetics. 39 (12): 1443–1452. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549. Retrieved 21 February 2022.
Menon IA, Basu PK, Persad S, Avaria M, Felix CC, Kalyanaraman B (1987). "Is there any difference in the photobiological properties of melanins isolated from human blue and brown eyes?". Br J Ophthalmol. 71 (7): 549–52. doi:10.1136/bjo.71.7.549. PMC 1041224. PMID 2820463.
Eiberg, Hans; Troelsen, Jesper; Nielsen, Mette; Mikkelsen, Annemette; Mengel-From, Jonas; Kjaer, Klaus W.; Hansen, Lars (2008). "Blue eye color in humans may be caused by a perfectly associated founder mutation in a regulatory element located within the HERC2 gene inhibiting OCA2 expression". Hum. Genet. 123 (2): 177–87. doi:10.1007/s00439-007-0460-x. PMID 18172690. S2CID 9886658.
Lucock, Mark D. (25 June 2022). "The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and <scp>UV</scp> radiation during human expansion". American Journal of Biological Anthropology. 180 (2). Wiley: 252–271. doi:10.1002/ajpa.24564. ISSN 2692-7691. PMC 10083917. PMID 36790744. This may also be a direct counter‐measure to a short winter photoperiod at high latitudes; blue eyes increase intraocular light scattering and thereby suppress melatonin release from the pineal gland. This may be an adaptive trait to reduce/prevent depression, a condition linked to short day length (Higuchi et al., 2007; Lucock et al., 2021).
Kastelic, V; Pośpiech, E; Draus-Barini, J; Branicki, W; Drobnič, K (2013). "Prediction of eye color in the Slovenian population using the IrisPlex SNPs". Croat. Med. J. 54 (4): 381–6. doi:10.3325/cmj.2013.54.381. PMC 3760663. PMID 23986280.
Fertl, Dagmar; Rosel, Patricia (2009). "Albinism". Albinism. Encyclopedia of Marine Mammals (Second ed.). Academic Press. pp. 24–26. doi:10.1016/b978-0-12-373553-9.00006-7. ISBN 978-0-12-373553-9. Retrieved 11 October 2022.
White, Désirée; Rabago-Smith, Montserrat (2010). "Genotype–phenotype associations and human eye color". Journal of Human Genetics. 56 (1): 5–7. doi:10.1038/jhg.2010.126. PMID 20944644. S2CID 33808164.
Solano, F. (2014). "Melanins: Skin Pigments and Much More—Types, Structural Models, Biological Functions, and Formation Routes". New Journal of Science. 2014. hindawi.com: 1–28. doi:10.1155/2014/498276. Eye color change and the affecting factors.
Nicolas, Caroline M; Robman, Luba D; Tikellis, Gabriella; Dimitrov, Peter N; Dowrick, Adam; Guymer, Robyn H; McCarty, Catherine A (2003). "Iris colour, ethnic origin and progression of age-related macular degeneration". Clin. Experiment. Ophthalmol. 31 (6): 465–9. doi:10.1046/j.1442-9071.2003.00711.x. PMID 14641151. S2CID 25878963.
Rafnsson V, Hrafnkelsson J, Tulinius H, Sigurgeirsson B, Olafsson JH (2004). "Risk factors for malignant melanoma in an Icelandic population sample". Prev Med. 39 (2): 247–52. doi:10.1016/j.ypmed.2004.03.027. PMID 15226032.
Stang A, Ahrens W, Anastassiou G, Jöckel KH (2003). "Phenotypical characteristics, lifestyle, social class and uveal melanoma". Ophthalmic Epidemiol. 10 (5): 293–302. doi:10.1076/opep.10.5.293.17319. PMID 14566630. S2CID 1592701.
Cumming RG, Mitchell P, Lim R (2000). "Iris color and cataract: The Blue Mountains Eye Study". American Journal of Ophthalmology. 130 (2): 237–238. doi:10.1016/S0002-9394(00)00479-7. PMID 11004303.
McDonnell G, Esmonde T (1999). "A homesick student". Postgrad Med J. 75 (884): 375–8. doi:10.1136/pgmj.75.884.375. PMC 1741256. PMID 10435182.
de la Maza, Maite Sainz; Tauber, Joseph; Foster, C. Stephen (2012). "Noninflammatory Diseases of the Sclera". The Sclera. pp. 277–297. doi:10.1007/978-1-4419-6502-8_8. ISBN 978-1-4419-6501-1.
Wheatley, TJ (2006). "Upper gastrointestinal surgery". In Kingsnorth, Andrew N; Majid, Aljafri A (eds.). Fundamentals of Surgical Practice. pp. 230–248. doi:10.1017/CBO9780511545740.013. ISBN 978-0-511-54574-0.
Imesch PD, Wallow IH, Albert DM (1997). "The color of the human eye: a review of morphologic correlates and of some conditions that affect iridial pigmentation". Surv Ophthalmol. 41 (Suppl 2): S117–23. doi:10.1016/S0039-6257(97)80018-5. PMID 9154287.
Hejkal TW, Camras CB (1999). "Prostaglandin analogs in the treatment of glaucoma". Seminars in Ophthalmology. 14 (3): 114–23. doi:10.3109/08820539909061464. PMID 10790575.
Darren Peshek; Negar Semmaknejad; Donald Hoffman; Pete Foley (2011). "Preliminary evidence that the limbal ring influences facial attractiveness" (PDF). Evolutionary Psychology. 9 (2): 137–146. doi:10.1177/147470491100900201. PMC 10519137. PMID 22947961.
Brown M, Sacco DF (2018). "Put a (Limbal) Ring on It: Women Perceive Men's Limbal Rings as a Health Cue in Short-Term Mating Domains". Pers Soc Psychol Bull. 44 (1): 80–91. doi:10.1177/0146167217733072. PMID 28978250.
Miller, L. K.; Rowe, P. J.; Lund, J. (August 1992). "Correlation of eye color on self-paced and reactive motor performance". Perceptual and Motor Skills. 75 (1): 91–95. doi:10.2466/pms.1992.75.1.91. ISSN 0031-5125. PMID 1528697. S2CID 31398375.
Rowe, P. J.; Evans, P. (August 1994). "Ball color, eye color, and a reactive motor skill". Perceptual and Motor Skills. 79 (1 Pt 2): 671–674. doi:10.2466/pms.1994.79.1.671. ISSN 0031-5125. PMID 7808908. S2CID 6743916.
Crowe, M.; O'Connor, D. (October 2001). "Eye colour and reaction time to visual stimuli in rugby league players". Perceptual and Motor Skills. 93 (2): 455–460. doi:10.2466/pms.2001.93.2.455. ISSN 0031-5125. PMID 11769902. S2CID 22785194.
German EJ, Hurst MA, Wood D, Gilchrist J (1998). "A novel system for the objective classification of iris colour and its correlation with response to 1% tropicamide". Ophthalmic Physiol Opt. 18 (2): 103–10. doi:10.1016/S0275-5408(97)00070-7. PMID 9692029.
Piquet-Thepot, M.-M. (26 March 1968). "Contribution à l'anthropologie des Corses: Anthropologie de la tête (suite)". Bulletins et Mémoires de la Société d'anthropologie de Paris. 3 (3): 183–218. doi:10.3406/bmsap.1968.1417.

edromanguitars.com

Color Perception Archived 20 October 2006 at the Wayback Machine. Edromanguitars.com. Retrieved on 23 December 2011.

emedicine.com

Ocular Manifestations of Albinism at eMedicine

escholarship.org

Lefohn A, Budge B, Shirley P, Caruso R, Reinhard E (2003). "An Ocularist's Approach to Human Iris Synthesis". IEEE Comput. Graph. Appl. 23 (6): 70–5. doi:10.1109/MCG.2003.1242384. S2CID 537404.
Darren Peshek; Negar Semmaknejad; Donald Hoffman; Pete Foley (2011). "Preliminary evidence that the limbal ring influences facial attractiveness" (PDF). Evolutionary Psychology. 9 (2): 137–146. doi:10.1177/147470491100900201. PMC 10519137. PMID 22947961.

eyecarecontacts.com

Larry Bickford Eye Color Archived 23 October 2010 at the Wayback Machine. Eyecarecontacts.com. Retrieved on 23 December 2011.

eyedoctorguide.com

Blue Eyes Versus Brown Eyes: A Primer on Eye Color Archived 17 October 2008 at the Wayback Machine. Eyedoctorguide.com. Retrieved on 23 December 2011.

handle.net

hdl.handle.net

Martinez-Cadenas, Conrado; Peña-Chilet, Maria; Ibarrola-Villava, Maider; Ribas, Gloria (2013). "Gender is a major factor explaining discrepancies in eye colour prediction based on HERC2/OCA2 genotype and the IrisPlex model". Forensic Science International: Genetics. 7 (4). Elsevier BV: 453–460. doi:10.1016/j.fsigen.2013.03.007. hdl:10234/86709. ISSN 1872-4973. PMID 23601698. "This effect is what may explain the fact that there seem to be comparatively higher frequencies of blue-eyed males than blue-eyed females in populations of European origin such as Iceland [10], Holland [10], Australia [18] or Poland [17], as well as in this study (see Fig. 2)."
Hernando, Barbara; Ibarrola-Villava, Maider; Fernandez, Lara P.; Peña-Chilet, Maria; Llorca-Cardeñosa, Marta; Oltra, Sara S.; Alonso, Santos; Boyano, Maria Dolores; Martinez-Cadenas, Conrado; Ribas, Gloria (2016). "Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin". Biology of Sex Differences. 7 (1): 17. doi:10.1186/s13293-016-0070-1. hdl:10810/32358. ISSN 2042-6410. PMC 4797181. PMID 26998216. The results of this study suggest that there are indeed sex-specific genetic effects in human pigmentation, with larger effects for darker pigmentation in females compared to males. A plausible cause might be the differentially expressed melanogenic genes in females due to higher oestrogen levels. These sex-specific genetic effects would help explain the presence of darker eye and skin pigmentation in females, as well as the well-known higher melanoma risk displayed by males.

harvard.edu

ui.adsabs.harvard.edu

Liu, Fan; Van Duijn, Kate; Vingerling, Johannes R.; Hofman, Albert; Uitterlinden, André G.; Janssens, A. Cecile J.W.; Kayser, Manfred (2009). "Eye color and the prediction of complex phenotypes from genotypes". Current Biology. 19 (5): R192 – R193. Bibcode:2009CBio...19.R192L. doi:10.1016/j.cub.2009.01.027. PMID 19278628.
Ruiz-Linares, Andrés; Adhikari, Kaustubh; Acuña-Alonzo, Victor; Quinto-Sanchez, Mirsha; Jaramillo, Claudia; Arias, William; Fuentes, Macarena; Pizarro, María; Everardo, Paola; de Avila, Francisco; Gómez-Valdés, Jorge; León-Mimila, Paola; Hunemeier, Tábita; Ramallo, Virginia; Silva de Cerqueira, Caio C. (25 September 2014). "Admixture in Latin America: Geographic Structure, Phenotypic Diversity and Self-Perception of Ancestry Based on 7,342 Individuals". PLOS Genetics. 10 (9): e1004572. Bibcode:2014PLOSG..10.4572R. doi:10.1371/journal.pgen.1004572. ISSN 1553-7390. PMC 4177621. PMID 25254375.

hhmi.org

Howard Hughes Medical Institute: Ask A Scientist Archived 1 September 2010 at the Wayback Machine. Hhmi.org. Retrieved on 23 December 2011.

imdb.com

Biography for Lee Van Cleef at IMDb

independent.co.uk

"How one ancestor helped turn our brown eyes blue". The Independent. 31 January 2008. Retrieved 21 December 2015. "Everyone with blue eyes alive today can trace their ancestry back to one person who probably lived about 10,000 years ago in the Black Sea region, a study has found."

iovs.org

Seddon, JM; CR Sahagian; RJ Glynn; RD Sperduto; ES Gragoudas (1 August 1990). "Evaluation of an iris color classification system". Investigative Ophthalmology & Visual Science. 31 (8): 1592–8. PMID 2201662. Retrieved 19 October 2011.

medscape.com

emedicine.medscape.com

Aniridia at eMedicine

medterms.com

"Eye color definition – Medical Dictionary definitions of popular medical terms easily defined on MedTerms". Medterms.com. 29 October 2003. Archived from the original on 5 June 2011. Retrieved 19 October 2011.

nbcnews.com

Bryner, Jeanna (31 January 2008). "Genetic mutation makes those brown eyes blue". NBC News. Archived from the original on 1 July 2013. Retrieved 19 October 2009.

newscientist.com

"DNA test for eye colour could help fight crime", New Scientist 14 March 2009.

nih.gov

pubmed.ncbi.nlm.nih.gov

Wielgus AR, Sarna T (2005). "Melanin in human irides of different color and age of donors". Pigment Cell Res. 18 (6): 454–64. doi:10.1111/j.1600-0749.2005.00268.x. ISSN 0893-5785. PMID 16280011.
Prota G, Hu DN, Vincensi MR, McCormick SA, Napolitano A (1998). "Characterization of melanins in human irides and cultured uveal melanocytes from eyes of different colors". Exp. Eye Res. 67 (3): 293–9. doi:10.1006/exer.1998.0518. PMID 9778410.
Oliphant LW (1987). "Pteridines and purines as major pigments of the avian iris". Pigment Cell Res. 1 (2): 129–31. doi:10.1111/j.1600-0749.1987.tb00401.x. PMID 3507666.
Kayser, Manfred; Liu, Fan; Janssens, A. Cecile J.W.; Rivadeneira, Fernando; Lao, Oscar; Van Duijn, Kate; Vermeulen, Mark; Arp, Pascal; et al. (2008). "Three genome-wide association studies and a linkage analysis identify HERC2 as a human iris color gene". Am. J. Hum. Genet. 82 (2): 411–23. doi:10.1016/j.ajhg.2007.10.003. PMC 2427174. PMID 18252221.
White, Désirée; Rabago-Smith, Montserrat (14 October 2010). "Genotype–phenotype associations and human eye color". Journal of Human Genetics. 56 (1): 5–7. doi:10.1038/jhg.2010.126. PMID 20944644.
Duffy, David L.; Montgomery, Grant W.; Chen, Wei; Zhao, Zhen Zhen; Le, Lien; James, Michael R.; Hayward, Nicholas K.; Martin, Nicholas G.; Sturm, Richard A. (2007). "A three-single-nucleotide polymorphism haplotype in intron 1 of OCA2 explains most human eye-color variation". Am. J. Hum. Genet. 80 (2): 241–52. doi:10.1086/510885. PMC 1785344. PMID 17236130.
Sturm RA, Frudakis TN (2004). "Eye colour: portals into pigmentation genes and ancestry" (PDF). Trends Genet. 20 (8): 327–32. doi:10.1016/j.tig.2004.06.010. PMID 15262401. Archived from the original (PDF) on 9 September 2006.
Grant MD, Lauderdale DS (2002). "Cohort effects in a genetically determined trait: eye colour among US whites". Ann. Hum. Biol. 29 (6): 657–66. doi:10.1080/03014460210157394. PMID 12573082. S2CID 25364754.
Liu, Fan; Van Duijn, Kate; Vingerling, Johannes R.; Hofman, Albert; Uitterlinden, André G.; Janssens, A. Cecile J.W.; Kayser, Manfred (2009). "Eye color and the prediction of complex phenotypes from genotypes". Current Biology. 19 (5): R192 – R193. Bibcode:2009CBio...19.R192L. doi:10.1016/j.cub.2009.01.027. PMID 19278628.
Martinez-Cadenas, Conrado; Peña-Chilet, Maria; Ibarrola-Villava, Maider; Ribas, Gloria (2013). "Gender is a major factor explaining discrepancies in eye colour prediction based on HERC2/OCA2 genotype and the IrisPlex model". Forensic Science International: Genetics. 7 (4). Elsevier BV: 453–460. doi:10.1016/j.fsigen.2013.03.007. hdl:10234/86709. ISSN 1872-4973. PMID 23601698. "This effect is what may explain the fact that there seem to be comparatively higher frequencies of blue-eyed males than blue-eyed females in populations of European origin such as Iceland [10], Holland [10], Australia [18] or Poland [17], as well as in this study (see Fig. 2)."
Hernando, Barbara; Ibarrola-Villava, Maider; Fernandez, Lara P.; Peña-Chilet, Maria; Llorca-Cardeñosa, Marta; Oltra, Sara S.; Alonso, Santos; Boyano, Maria Dolores; Martinez-Cadenas, Conrado; Ribas, Gloria (2016). "Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin". Biology of Sex Differences. 7 (1): 17. doi:10.1186/s13293-016-0070-1. hdl:10810/32358. ISSN 2042-6410. PMC 4797181. PMID 26998216. The results of this study suggest that there are indeed sex-specific genetic effects in human pigmentation, with larger effects for darker pigmentation in females compared to males. A plausible cause might be the differentially expressed melanogenic genes in females due to higher oestrogen levels. These sex-specific genetic effects would help explain the presence of darker eye and skin pigmentation in females, as well as the well-known higher melanoma risk displayed by males.
Hanel, Andrea; Carlberg, Carsten (September 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 864–875. doi:10.1111/exd.14142. ISSN 0906-6705. PMID 32621306. S2CID 220335539. "The genetic history of today's European populations is based on continuous migrations over the past 40 000 years. Homo sapiens arrived in Europe from Near East some 42 000 years ago.[48, 49] Like in their African origin, these humans had dark skin but due to variations of their OCA2 gene (causing iris depigmentation) many of them had blue eyes[48,50] (Figure 2, left)." Figure 2: "Phenotype information was retrieved from supplementary files of the according publications or additionally assessed following the instructions of HirisPlex-S, which is a forensic DNA phenotyping tool based on an array of different marker SNPs providing additional support to the SNPs of the genes SLC24A5, SLC45A2 and OCA2."
Sulem, Patrick; Gudbjartsson, Daniel F; Stacey, Simon N; Helgason, Agnar; Rafnar, Thorunn; Magnusson, Kristinn P; Manolescu, Andrei; Karason, Ari; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nat. Genet. 39 (12): 1443–52. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549.
Liu, Fan; Wollstein, Andreas; Hysi, Pirro G.; Ankra-Badu, Georgina A.; Spector, Timothy D.; Park, Daniel; Zhu, Gu; Larsson, Mats; Duffy, David L.; Montgomery, Grant W.; MacKey, David A.; Walsh, Susan; Lao, Oscar; Hofman, Albert; Rivadeneira, Fernando; Vingerling, Johannes R.; Uitterlinden, André G.; Martin, Nicholas G.; Hammond, Christopher J.; Kayser, Manfred (2010). "Digital Quantification of Human Eye Color Highlights Genetic Association of Three New Loci". PLOS Genetics. 6 (5): e1000934. doi:10.1371/journal.pgen.1000934. PMC 2865509. PMID 20463881.
Bito, LZ (1997). "Eye Color Changes Past Early Childhood". Archives of Ophthalmology. 115 (5): 659–63. doi:10.1001/archopht.1997.01100150661017. PMID 9152135.
Brenda J Bradley, Anja Pedersen, Nicholas I Mundy: Blue eyes in lemurs and humans: same phenotype, different genetic mechanism Am J Phys Anthropol. 2009
Oliphant LW (1987). "Observations on the pigmentation of the pigeon iris". Pigment Cell Res. 1 (3): 202–8. doi:10.1111/j.1600-0749.1987.tb00414.x. PMID 3508278.
Oliphant LW (1981). "Crystalline pteridines in the stromal pigment cells of the iris of the great horned owl". Cell Tissue Res. 217 (2): 387–95. doi:10.1007/BF00233588. PMID 7237534. S2CID 8061493.
Ruiz-Linares, Andrés; Adhikari, Kaustubh; Acuña-Alonzo, Victor; Quinto-Sanchez, Mirsha; Jaramillo, Claudia; Arias, William; Fuentes, Macarena; Pizarro, María; Everardo, Paola; de Avila, Francisco; Gómez-Valdés, Jorge; León-Mimila, Paola; Hunemeier, Tábita; Ramallo, Virginia; Silva de Cerqueira, Caio C. (25 September 2014). "Admixture in Latin America: Geographic Structure, Phenotypic Diversity and Self-Perception of Ancestry Based on 7,342 Individuals". PLOS Genetics. 10 (9): e1004572. Bibcode:2014PLOSG..10.4572R. doi:10.1371/journal.pgen.1004572. ISSN 1553-7390. PMC 4177621. PMID 25254375.
Zhu, Gu; Evans, David M.; Duffy, David L.; Montgomery, Grant W.; Medland, Sarah E.; Gillespie, Nathan A.; Ewen, Kelly R.; Jewell, Mary; Liew, Yew Wah; Hayward, Nicholas K.; Sturma, Richard A.; Trenta, Jeffrey M.; Martina, Nicholas G. (2004). "A genome scan for eye color in 502 twin families: most variation is due to a QTL on chromosome 15q". Twin Res. 7 (2): 197–210. doi:10.1375/136905204323016186. PMID 15169604.
Albert, Daniel M; Green, W Richard; Zimbric, Michele L; Lo, Cecilia; Gangnon, Ronald E; Hope, Kirsten L; Gleiser, Joel (2003). "Iris melanocyte numbers in Asian, African American, and Caucasian irides". Transactions of the American Ophthalmological Society. 101: 217–222. PMC 1358991. PMID 14971580.
Mitchell R, Rochtchina E, Lee A, Wang JJ, Mitchell P (2003). "Iris color and intraocular pressure: the Blue Mountains Eye Study". Am. J. Ophthalmol. 135 (3): 384–6. doi:10.1016/S0002-9394(02)01967-0. PMID 12614760.
Lindsey JD, Jones HL, Hewitt EG, Angert M, Weinreb RN (2001). "Induction of tyrosinase gene transcription in human iris organ cultures exposed to latanoprost". Arch. Ophthalmol. 119 (6): 853–60. doi:10.1001/archopht.119.6.853. PMID 11405836.
Frank RN, Puklin JE, Stock C, Canter LA (2000). "Race, iris color, and age-related macular degeneration". Trans Am Ophthalmol Soc. 98: 109–15, discussion 115–7. PMC 1298217. PMID 11190014.
Regan S, Judge HE, Gragoudas ES, Egan KM (1999). "Iris color as a prognostic factor in ocular melanoma". Arch. Ophthalmol. 117 (6): 811–4. doi:10.1001/archopht.117.6.811. PMID 10369595.
Hawkins TA, Stewart WC, McMillan TA, Gwynn DR (1994). "Analysis of diode, argon, and Nd: YAG peripheral iridectomy in cadaver eyes". Doc Ophthalmol. 87 (4): 367–76. doi:10.1007/BF01203345. PMID 7851220. S2CID 30893783.
Hammond BR, Fuld K, Snodderly DM (1996). "Iris color and macular pigment optical density". Exp. Eye Res. 62 (3): 293–7. doi:10.1006/exer.1996.0035. PMID 8690039.
Muiños Díaz, Yerena; Saornil, Maria A.; Almaraz, Ana; Muñoz-Moreno, Mari F.; García, Ciro; Sanz, Ruperto (2009). "Iris color: validation of a new classification and distribution in a Spanish population-based sample". European Journal of Ophthalmology. 19 (4): 686–689. doi:10.1177/112067210901900427. ISSN 1120-6721. PMID 19551689. S2CID 40940828.
Keyser, Christine; Bouakaze, Caroline; Crubézy, Eric; Nikolaev, Valery G.; Montagnon, Daniel; Reis, Tatiana; Ludes, Bertrand (2009). "Ancient DNA Provides New Insights into the History of South Siberian Kurgan People". Human Genetics. 126 (3): 395–410. doi:10.1007/s00439-009-0683-0. PMID 19449030. S2CID 21347353. Indeed, among the SNPs tested was rs12913832, a single DNA variation within a regulatory element of HERC2 gene which is associated to blue eye color in humans. This polymorphism, together with the diplotypes obtained from variations of the OCA2 locus (major contributor to the human eye color variation) showed that at least 60% of the ancient Siberian specimens under study had blue (or green) eyes.
Sulem, P.; Gudbjartsson, D.; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nature Genetics. 39 (12): 1443–1452. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549. Retrieved 21 February 2022.
Menon IA, Basu PK, Persad S, Avaria M, Felix CC, Kalyanaraman B (1987). "Is there any difference in the photobiological properties of melanins isolated from human blue and brown eyes?". Br J Ophthalmol. 71 (7): 549–52. doi:10.1136/bjo.71.7.549. PMC 1041224. PMID 2820463.
Eiberg, Hans; Troelsen, Jesper; Nielsen, Mette; Mikkelsen, Annemette; Mengel-From, Jonas; Kjaer, Klaus W.; Hansen, Lars (2008). "Blue eye color in humans may be caused by a perfectly associated founder mutation in a regulatory element located within the HERC2 gene inhibiting OCA2 expression". Hum. Genet. 123 (2): 177–87. doi:10.1007/s00439-007-0460-x. PMID 18172690. S2CID 9886658.
Lucock, Mark D. (25 June 2022). "The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and <scp>UV</scp> radiation during human expansion". American Journal of Biological Anthropology. 180 (2). Wiley: 252–271. doi:10.1002/ajpa.24564. ISSN 2692-7691. PMC 10083917. PMID 36790744. This may also be a direct counter‐measure to a short winter photoperiod at high latitudes; blue eyes increase intraocular light scattering and thereby suppress melatonin release from the pineal gland. This may be an adaptive trait to reduce/prevent depression, a condition linked to short day length (Higuchi et al., 2007; Lucock et al., 2021).
Kastelic, V; Pośpiech, E; Draus-Barini, J; Branicki, W; Drobnič, K (2013). "Prediction of eye color in the Slovenian population using the IrisPlex SNPs". Croat. Med. J. 54 (4): 381–6. doi:10.3325/cmj.2013.54.381. PMC 3760663. PMID 23986280.
White, Désirée; Rabago-Smith, Montserrat (2010). "Genotype–phenotype associations and human eye color". Journal of Human Genetics. 56 (1): 5–7. doi:10.1038/jhg.2010.126. PMID 20944644. S2CID 33808164.
Nicolas, Caroline M; Robman, Luba D; Tikellis, Gabriella; Dimitrov, Peter N; Dowrick, Adam; Guymer, Robyn H; McCarty, Catherine A (2003). "Iris colour, ethnic origin and progression of age-related macular degeneration". Clin. Experiment. Ophthalmol. 31 (6): 465–9. doi:10.1046/j.1442-9071.2003.00711.x. PMID 14641151. S2CID 25878963.
Rafnsson V, Hrafnkelsson J, Tulinius H, Sigurgeirsson B, Olafsson JH (2004). "Risk factors for malignant melanoma in an Icelandic population sample". Prev Med. 39 (2): 247–52. doi:10.1016/j.ypmed.2004.03.027. PMID 15226032.
Stang A, Ahrens W, Anastassiou G, Jöckel KH (2003). "Phenotypical characteristics, lifestyle, social class and uveal melanoma". Ophthalmic Epidemiol. 10 (5): 293–302. doi:10.1076/opep.10.5.293.17319. PMID 14566630. S2CID 1592701.
Cumming RG, Mitchell P, Lim R (2000). "Iris color and cataract: The Blue Mountains Eye Study". American Journal of Ophthalmology. 130 (2): 237–238. doi:10.1016/S0002-9394(00)00479-7. PMID 11004303.
McDonnell G, Esmonde T (1999). "A homesick student". Postgrad Med J. 75 (884): 375–8. doi:10.1136/pgmj.75.884.375. PMC 1741256. PMID 10435182.
Imesch PD, Wallow IH, Albert DM (1997). "The color of the human eye: a review of morphologic correlates and of some conditions that affect iridial pigmentation". Surv Ophthalmol. 41 (Suppl 2): S117–23. doi:10.1016/S0039-6257(97)80018-5. PMID 9154287.
Hejkal TW, Camras CB (1999). "Prostaglandin analogs in the treatment of glaucoma". Seminars in Ophthalmology. 14 (3): 114–23. doi:10.3109/08820539909061464. PMID 10790575.
Darren Peshek; Negar Semmaknejad; Donald Hoffman; Pete Foley (2011). "Preliminary evidence that the limbal ring influences facial attractiveness" (PDF). Evolutionary Psychology. 9 (2): 137–146. doi:10.1177/147470491100900201. PMC 10519137. PMID 22947961.
Brown M, Sacco DF (2018). "Put a (Limbal) Ring on It: Women Perceive Men's Limbal Rings as a Health Cue in Short-Term Mating Domains". Pers Soc Psychol Bull. 44 (1): 80–91. doi:10.1177/0146167217733072. PMID 28978250.
Miller, L. K.; Rowe, P. J.; Lund, J. (August 1992). "Correlation of eye color on self-paced and reactive motor performance". Perceptual and Motor Skills. 75 (1): 91–95. doi:10.2466/pms.1992.75.1.91. ISSN 0031-5125. PMID 1528697. S2CID 31398375.
Rowe, P. J.; Evans, P. (August 1994). "Ball color, eye color, and a reactive motor skill". Perceptual and Motor Skills. 79 (1 Pt 2): 671–674. doi:10.2466/pms.1994.79.1.671. ISSN 0031-5125. PMID 7808908. S2CID 6743916.
Crowe, M.; O'Connor, D. (October 2001). "Eye colour and reaction time to visual stimuli in rugby league players". Perceptual and Motor Skills. 93 (2): 455–460. doi:10.2466/pms.2001.93.2.455. ISSN 0031-5125. PMID 11769902. S2CID 22785194.
German EJ, Hurst MA, Wood D, Gilchrist J (1998). "A novel system for the objective classification of iris colour and its correlation with response to 1% tropicamide". Ophthalmic Physiol Opt. 18 (2): 103–10. doi:10.1016/S0275-5408(97)00070-7. PMID 9692029.
Seddon, JM; CR Sahagian; RJ Glynn; RD Sperduto; ES Gragoudas (1 August 1990). "Evaluation of an iris color classification system". Investigative Ophthalmology & Visual Science. 31 (8): 1592–8. PMID 2201662. Retrieved 19 October 2011.

ncbi.nlm.nih.gov

Kayser, Manfred; Liu, Fan; Janssens, A. Cecile J.W.; Rivadeneira, Fernando; Lao, Oscar; Van Duijn, Kate; Vermeulen, Mark; Arp, Pascal; et al. (2008). "Three genome-wide association studies and a linkage analysis identify HERC2 as a human iris color gene". Am. J. Hum. Genet. 82 (2): 411–23. doi:10.1016/j.ajhg.2007.10.003. PMC 2427174. PMID 18252221.
Duffy, David L.; Montgomery, Grant W.; Chen, Wei; Zhao, Zhen Zhen; Le, Lien; James, Michael R.; Hayward, Nicholas K.; Martin, Nicholas G.; Sturm, Richard A. (2007). "A three-single-nucleotide polymorphism haplotype in intron 1 of OCA2 explains most human eye-color variation". Am. J. Hum. Genet. 80 (2): 241–52. doi:10.1086/510885. PMC 1785344. PMID 17236130.
Hernando, Barbara; Ibarrola-Villava, Maider; Fernandez, Lara P.; Peña-Chilet, Maria; Llorca-Cardeñosa, Marta; Oltra, Sara S.; Alonso, Santos; Boyano, Maria Dolores; Martinez-Cadenas, Conrado; Ribas, Gloria (2016). "Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin". Biology of Sex Differences. 7 (1): 17. doi:10.1186/s13293-016-0070-1. hdl:10810/32358. ISSN 2042-6410. PMC 4797181. PMID 26998216. The results of this study suggest that there are indeed sex-specific genetic effects in human pigmentation, with larger effects for darker pigmentation in females compared to males. A plausible cause might be the differentially expressed melanogenic genes in females due to higher oestrogen levels. These sex-specific genetic effects would help explain the presence of darker eye and skin pigmentation in females, as well as the well-known higher melanoma risk displayed by males.
Liu, Fan; Wollstein, Andreas; Hysi, Pirro G.; Ankra-Badu, Georgina A.; Spector, Timothy D.; Park, Daniel; Zhu, Gu; Larsson, Mats; Duffy, David L.; Montgomery, Grant W.; MacKey, David A.; Walsh, Susan; Lao, Oscar; Hofman, Albert; Rivadeneira, Fernando; Vingerling, Johannes R.; Uitterlinden, André G.; Martin, Nicholas G.; Hammond, Christopher J.; Kayser, Manfred (2010). "Digital Quantification of Human Eye Color Highlights Genetic Association of Three New Loci". PLOS Genetics. 6 (5): e1000934. doi:10.1371/journal.pgen.1000934. PMC 2865509. PMID 20463881.
Ruiz-Linares, Andrés; Adhikari, Kaustubh; Acuña-Alonzo, Victor; Quinto-Sanchez, Mirsha; Jaramillo, Claudia; Arias, William; Fuentes, Macarena; Pizarro, María; Everardo, Paola; de Avila, Francisco; Gómez-Valdés, Jorge; León-Mimila, Paola; Hunemeier, Tábita; Ramallo, Virginia; Silva de Cerqueira, Caio C. (25 September 2014). "Admixture in Latin America: Geographic Structure, Phenotypic Diversity and Self-Perception of Ancestry Based on 7,342 Individuals". PLOS Genetics. 10 (9): e1004572. Bibcode:2014PLOSG..10.4572R. doi:10.1371/journal.pgen.1004572. ISSN 1553-7390. PMC 4177621. PMID 25254375.
Albert, Daniel M; Green, W Richard; Zimbric, Michele L; Lo, Cecilia; Gangnon, Ronald E; Hope, Kirsten L; Gleiser, Joel (2003). "Iris melanocyte numbers in Asian, African American, and Caucasian irides". Transactions of the American Ophthalmological Society. 101: 217–222. PMC 1358991. PMID 14971580.
Frank RN, Puklin JE, Stock C, Canter LA (2000). "Race, iris color, and age-related macular degeneration". Trans Am Ophthalmol Soc. 98: 109–15, discussion 115–7. PMC 1298217. PMID 11190014.
Menon IA, Basu PK, Persad S, Avaria M, Felix CC, Kalyanaraman B (1987). "Is there any difference in the photobiological properties of melanins isolated from human blue and brown eyes?". Br J Ophthalmol. 71 (7): 549–52. doi:10.1136/bjo.71.7.549. PMC 1041224. PMID 2820463.
Lucock, Mark D. (25 June 2022). "The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and <scp>UV</scp> radiation during human expansion". American Journal of Biological Anthropology. 180 (2). Wiley: 252–271. doi:10.1002/ajpa.24564. ISSN 2692-7691. PMC 10083917. PMID 36790744. This may also be a direct counter‐measure to a short winter photoperiod at high latitudes; blue eyes increase intraocular light scattering and thereby suppress melatonin release from the pineal gland. This may be an adaptive trait to reduce/prevent depression, a condition linked to short day length (Higuchi et al., 2007; Lucock et al., 2021).
Kastelic, V; Pośpiech, E; Draus-Barini, J; Branicki, W; Drobnič, K (2013). "Prediction of eye color in the Slovenian population using the IrisPlex SNPs". Croat. Med. J. 54 (4): 381–6. doi:10.3325/cmj.2013.54.381. PMC 3760663. PMID 23986280.
McDonnell G, Esmonde T (1999). "A homesick student". Postgrad Med J. 75 (884): 375–8. doi:10.1136/pgmj.75.884.375. PMC 1741256. PMID 10435182.
Darren Peshek; Negar Semmaknejad; Donald Hoffman; Pete Foley (2011). "Preliminary evidence that the limbal ring influences facial attractiveness" (PDF). Evolutionary Psychology. 9 (2): 137–146. doi:10.1177/147470491100900201. PMC 10519137. PMID 22947961.

nytimes.com

Belkin, Douglas (18 October 2006). "Blue eyes are increasingly rare in America - Americas - International Herald Tribune (Published 2006)". The New York Times. ISSN 0362-4331. Retrieved 2 February 2021.
O'Connor, Anahad (19 January 2009). "The Claim: Eye Color Can Have an Effect on Vision". The New York Times. ISSN 0362-4331. Retrieved 5 May 2021.

ohio-state.edu

biosci.ohio-state.edu

Sturm RA, Frudakis TN (2004). "Eye colour: portals into pigmentation genes and ancestry" (PDF). Trends Genet. 20 (8): 327–32. doi:10.1016/j.tig.2004.06.010. PMID 15262401. Archived from the original (PDF) on 9 September 2006.

omim.org

Online Mendelian Inheritance in Man (OMIM): SKIN/HAIR/EYE PIGMENTATION, VARIATION IN, 1; SHEP1 - 227220

pcworld.co.nz

Dave Johnson (16 January 2009). "HOW TO: Avoid the red eye effect". New Zealand PC World. Archived from the original on 24 February 2010. Retrieved 9 January 2010.

persee.fr

Piquet-Thepot, M.-M. (26 March 1968). "Contribution à l'anthropologie des Corses: Anthropologie de la tête (suite)". Bulletins et Mémoires de la Société d'anthropologie de Paris. 3 (3): 183–218. doi:10.3406/bmsap.1968.1417.

psu.edu

citeseerx.ist.psu.edu

Huiqiong Wang; Lin, S.; Xiaopei Liu; Sing Bing Kang (2005). "Separating reflections in human iris images for illumination estimation". Tenth IEEE International Conference on Computer Vision (ICCV'05) Volume 1. pp. 1691–1698 Vol. 2. CiteSeerX 10.1.1.87.418. doi:10.1109/ICCV.2005.215. ISBN 978-0-7695-2334-7. S2CID 2215768.

researchgate.net

Sulem, P.; Gudbjartsson, D.; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nature Genetics. 39 (12): 1443–1452. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549. Retrieved 21 February 2022.

sciencedaily.com

No Single Gene For Eye Color, Researchers Prove. Sciencedaily.com (22 February 2007). Retrieved on 2011-12-23.

sciencedirect.com

Fertl, Dagmar; Rosel, Patricia (2009). "Albinism". Albinism. Encyclopedia of Marine Mammals (Second ed.). Academic Press. pp. 24–26. doi:10.1016/b978-0-12-373553-9.00006-7. ISBN 978-0-12-373553-9. Retrieved 11 October 2022.

semanticscholar.org

api.semanticscholar.org

Huiqiong Wang; Lin, S.; Xiaopei Liu; Sing Bing Kang (2005). "Separating reflections in human iris images for illumination estimation". Tenth IEEE International Conference on Computer Vision (ICCV'05) Volume 1. pp. 1691–1698 Vol. 2. CiteSeerX 10.1.1.87.418. doi:10.1109/ICCV.2005.215. ISBN 978-0-7695-2334-7. S2CID 2215768.
Grant MD, Lauderdale DS (2002). "Cohort effects in a genetically determined trait: eye colour among US whites". Ann. Hum. Biol. 29 (6): 657–66. doi:10.1080/03014460210157394. PMID 12573082. S2CID 25364754.
Hanel, Andrea; Carlberg, Carsten (September 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 864–875. doi:10.1111/exd.14142. ISSN 0906-6705. PMID 32621306. S2CID 220335539. "The genetic history of today's European populations is based on continuous migrations over the past 40 000 years. Homo sapiens arrived in Europe from Near East some 42 000 years ago.[48, 49] Like in their African origin, these humans had dark skin but due to variations of their OCA2 gene (causing iris depigmentation) many of them had blue eyes[48,50] (Figure 2, left)." Figure 2: "Phenotype information was retrieved from supplementary files of the according publications or additionally assessed following the instructions of HirisPlex-S, which is a forensic DNA phenotyping tool based on an array of different marker SNPs providing additional support to the SNPs of the genes SLC24A5, SLC45A2 and OCA2."
Sulem, Patrick; Gudbjartsson, Daniel F; Stacey, Simon N; Helgason, Agnar; Rafnar, Thorunn; Magnusson, Kristinn P; Manolescu, Andrei; Karason, Ari; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nat. Genet. 39 (12): 1443–52. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549.
Oliphant LW (1981). "Crystalline pteridines in the stromal pigment cells of the iris of the great horned owl". Cell Tissue Res. 217 (2): 387–95. doi:10.1007/BF00233588. PMID 7237534. S2CID 8061493.
Lefohn A, Budge B, Shirley P, Caruso R, Reinhard E (2003). "An Ocularist's Approach to Human Iris Synthesis". IEEE Comput. Graph. Appl. 23 (6): 70–5. doi:10.1109/MCG.2003.1242384. S2CID 537404.
Hawkins TA, Stewart WC, McMillan TA, Gwynn DR (1994). "Analysis of diode, argon, and Nd: YAG peripheral iridectomy in cadaver eyes". Doc Ophthalmol. 87 (4): 367–76. doi:10.1007/BF01203345. PMID 7851220. S2CID 30893783.
Muiños Díaz, Yerena; Saornil, Maria A.; Almaraz, Ana; Muñoz-Moreno, Mari F.; García, Ciro; Sanz, Ruperto (2009). "Iris color: validation of a new classification and distribution in a Spanish population-based sample". European Journal of Ophthalmology. 19 (4): 686–689. doi:10.1177/112067210901900427. ISSN 1120-6721. PMID 19551689. S2CID 40940828.
Keyser, Christine; Bouakaze, Caroline; Crubézy, Eric; Nikolaev, Valery G.; Montagnon, Daniel; Reis, Tatiana; Ludes, Bertrand (2009). "Ancient DNA Provides New Insights into the History of South Siberian Kurgan People". Human Genetics. 126 (3): 395–410. doi:10.1007/s00439-009-0683-0. PMID 19449030. S2CID 21347353. Indeed, among the SNPs tested was rs12913832, a single DNA variation within a regulatory element of HERC2 gene which is associated to blue eye color in humans. This polymorphism, together with the diplotypes obtained from variations of the OCA2 locus (major contributor to the human eye color variation) showed that at least 60% of the ancient Siberian specimens under study had blue (or green) eyes.
Sulem, P.; Gudbjartsson, D.; et al. (2007). "Genetic determinants of hair, eye and skin pigmentation in Europeans". Nature Genetics. 39 (12): 1443–1452. doi:10.1038/ng.2007.13. PMID 17952075. S2CID 19313549. Retrieved 21 February 2022.
Eiberg, Hans; Troelsen, Jesper; Nielsen, Mette; Mikkelsen, Annemette; Mengel-From, Jonas; Kjaer, Klaus W.; Hansen, Lars (2008). "Blue eye color in humans may be caused by a perfectly associated founder mutation in a regulatory element located within the HERC2 gene inhibiting OCA2 expression". Hum. Genet. 123 (2): 177–87. doi:10.1007/s00439-007-0460-x. PMID 18172690. S2CID 9886658.
White, Désirée; Rabago-Smith, Montserrat (2010). "Genotype–phenotype associations and human eye color". Journal of Human Genetics. 56 (1): 5–7. doi:10.1038/jhg.2010.126. PMID 20944644. S2CID 33808164.
Nicolas, Caroline M; Robman, Luba D; Tikellis, Gabriella; Dimitrov, Peter N; Dowrick, Adam; Guymer, Robyn H; McCarty, Catherine A (2003). "Iris colour, ethnic origin and progression of age-related macular degeneration". Clin. Experiment. Ophthalmol. 31 (6): 465–9. doi:10.1046/j.1442-9071.2003.00711.x. PMID 14641151. S2CID 25878963.
Stang A, Ahrens W, Anastassiou G, Jöckel KH (2003). "Phenotypical characteristics, lifestyle, social class and uveal melanoma". Ophthalmic Epidemiol. 10 (5): 293–302. doi:10.1076/opep.10.5.293.17319. PMID 14566630. S2CID 1592701.
Miller, L. K.; Rowe, P. J.; Lund, J. (August 1992). "Correlation of eye color on self-paced and reactive motor performance". Perceptual and Motor Skills. 75 (1): 91–95. doi:10.2466/pms.1992.75.1.91. ISSN 0031-5125. PMID 1528697. S2CID 31398375.
Rowe, P. J.; Evans, P. (August 1994). "Ball color, eye color, and a reactive motor skill". Perceptual and Motor Skills. 79 (1 Pt 2): 671–674. doi:10.2466/pms.1994.79.1.671. ISSN 0031-5125. PMID 7808908. S2CID 6743916.
Crowe, M.; O'Connor, D. (October 2001). "Eye colour and reaction time to visual stimuli in rugby league players". Perceptual and Motor Skills. 93 (2): 455–460. doi:10.2466/pms.2001.93.2.455. ISSN 0031-5125. PMID 11769902. S2CID 22785194.

slate.com

Palmer, Roxanne (25 March 2005). "Elizabeth Taylor: Beautiful Mutant". Slate. Archived from the original on 27 March 2011. Retrieved 26 March 2011.

telegraph.co.uk

Highfield, Roger (30 January 2008). "Blue eyes result of ancient genetic 'mutation'". The Daily Telegraph. London. Archived from the original on 1 November 2009. Retrieved 19 October 2011.

thetech.org

Lucy Southworth. "Are gray eyes the same as blue in terms of genetics?". Understanding Genetics: Human Health and the Genome. Stanford School of Medicine. Archived from the original on 27 September 2011. Retrieved 19 October 2011.

tufts.edu

perseus.tufts.edu

Iliad 1:206 http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.01.0133%3Abook%3D1%3Acard%3D206

ucla.edu

cogweb.ucla.edu

Why Do Europeans Have So Many Hair and Eye Colors?. Cogweb.ucla.edu. Retrieved on 23 December 2011.

web.archive.org

"Eye color definition – Medical Dictionary definitions of popular medical terms easily defined on MedTerms". Medterms.com. 29 October 2003. Archived from the original on 5 June 2011. Retrieved 19 October 2011.
Sturm RA, Frudakis TN (2004). "Eye colour: portals into pigmentation genes and ancestry" (PDF). Trends Genet. 20 (8): 327–32. doi:10.1016/j.tig.2004.06.010. PMID 15262401. Archived from the original (PDF) on 9 September 2006.
Howard Hughes Medical Institute: Ask A Scientist Archived 1 September 2010 at the Wayback Machine. Hhmi.org. Retrieved on 23 December 2011.
Larry Bickford Eye Color Archived 23 October 2010 at the Wayback Machine. Eyecarecontacts.com. Retrieved on 23 December 2011.
Blue Eyes Versus Brown Eyes: A Primer on Eye Color Archived 17 October 2008 at the Wayback Machine. Eyedoctorguide.com. Retrieved on 23 December 2011.
"OCA2: The Gene for Color" Archived 6 October 2016 at the Wayback Machine. allaboutgenes.weebly.com. Retrieved on 8 September 2016.
Bryner, Jeanna (31 January 2008). "Genetic mutation makes those brown eyes blue". NBC News. Archived from the original on 1 July 2013. Retrieved 19 October 2009.
Highfield, Roger (30 January 2008). "Blue eyes result of ancient genetic 'mutation'". The Daily Telegraph. London. Archived from the original on 1 November 2009. Retrieved 19 October 2011.
"Distribution of Bodily Characters. Pigmentation, the Pilous System, and Morphology of the Soft Parts". Archived from the original on 26 July 2011.
Belkin, Douglas (17 October 2006). "Don't it make my blue eyes brown Americans are seeing a dramatic color change". The Boston Globe. Archived from the original on 22 October 2014.
Lucy Southworth. "Are gray eyes the same as blue in terms of genetics?". Understanding Genetics: Human Health and the Genome. Stanford School of Medicine. Archived from the original on 27 September 2011. Retrieved 19 October 2011.
NOAH – What is Albinism? Archived 14 May 2012 at the Wayback Machine. Albinism.org. Retrieved on 23 December 2011.
Dave Johnson (16 January 2009). "HOW TO: Avoid the red eye effect". New Zealand PC World. Archived from the original on 24 February 2010. Retrieved 9 January 2010.
Palmer, Roxanne (25 March 2005). "Elizabeth Taylor: Beautiful Mutant". Slate. Archived from the original on 27 March 2011. Retrieved 26 March 2011.
"Martin-Schulz Eye Color Chart". Archived from the original on 2 August 2016. Retrieved 13 January 2017.
Jones, S.L.; Schnirel, B.L. (2006). "Subspecies comparison of the Genus: Corucia". Polyphemos. 4 (1): 1–25. Archived from the original on 2 February 2009.
Color Perception Archived 20 October 2006 at the Wayback Machine. Edromanguitars.com. Retrieved on 23 December 2011.

weebly.com

allaboutgenes.weebly.com

"OCA2: The Gene for Color" Archived 6 October 2016 at the Wayback Machine. allaboutgenes.weebly.com. Retrieved on 8 September 2016.

wisc.edu

cs.wisc.edu

Fan S, Dyer CR, Hubbard L. Quantification and Correction of Iris Color." Technical report 1495, University of Wisconsin–Madison, Dec 2003.

worldatlas.com

"The World's Population By Eye Color". WorldAtlas. 6 October 2020. Retrieved 10 May 2021.

worldcat.org

search.worldcat.org

Wielgus AR, Sarna T (2005). "Melanin in human irides of different color and age of donors". Pigment Cell Res. 18 (6): 454–64. doi:10.1111/j.1600-0749.2005.00268.x. ISSN 0893-5785. PMID 16280011.
Martinez-Cadenas, Conrado; Peña-Chilet, Maria; Ibarrola-Villava, Maider; Ribas, Gloria (2013). "Gender is a major factor explaining discrepancies in eye colour prediction based on HERC2/OCA2 genotype and the IrisPlex model". Forensic Science International: Genetics. 7 (4). Elsevier BV: 453–460. doi:10.1016/j.fsigen.2013.03.007. hdl:10234/86709. ISSN 1872-4973. PMID 23601698. "This effect is what may explain the fact that there seem to be comparatively higher frequencies of blue-eyed males than blue-eyed females in populations of European origin such as Iceland [10], Holland [10], Australia [18] or Poland [17], as well as in this study (see Fig. 2)."
Hernando, Barbara; Ibarrola-Villava, Maider; Fernandez, Lara P.; Peña-Chilet, Maria; Llorca-Cardeñosa, Marta; Oltra, Sara S.; Alonso, Santos; Boyano, Maria Dolores; Martinez-Cadenas, Conrado; Ribas, Gloria (2016). "Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin". Biology of Sex Differences. 7 (1): 17. doi:10.1186/s13293-016-0070-1. hdl:10810/32358. ISSN 2042-6410. PMC 4797181. PMID 26998216. The results of this study suggest that there are indeed sex-specific genetic effects in human pigmentation, with larger effects for darker pigmentation in females compared to males. A plausible cause might be the differentially expressed melanogenic genes in females due to higher oestrogen levels. These sex-specific genetic effects would help explain the presence of darker eye and skin pigmentation in females, as well as the well-known higher melanoma risk displayed by males.
Hanel, Andrea; Carlberg, Carsten (September 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 864–875. doi:10.1111/exd.14142. ISSN 0906-6705. PMID 32621306. S2CID 220335539. "The genetic history of today's European populations is based on continuous migrations over the past 40 000 years. Homo sapiens arrived in Europe from Near East some 42 000 years ago.[48, 49] Like in their African origin, these humans had dark skin but due to variations of their OCA2 gene (causing iris depigmentation) many of them had blue eyes[48,50] (Figure 2, left)." Figure 2: "Phenotype information was retrieved from supplementary files of the according publications or additionally assessed following the instructions of HirisPlex-S, which is a forensic DNA phenotyping tool based on an array of different marker SNPs providing additional support to the SNPs of the genes SLC24A5, SLC45A2 and OCA2."
Ruiz-Linares, Andrés; Adhikari, Kaustubh; Acuña-Alonzo, Victor; Quinto-Sanchez, Mirsha; Jaramillo, Claudia; Arias, William; Fuentes, Macarena; Pizarro, María; Everardo, Paola; de Avila, Francisco; Gómez-Valdés, Jorge; León-Mimila, Paola; Hunemeier, Tábita; Ramallo, Virginia; Silva de Cerqueira, Caio C. (25 September 2014). "Admixture in Latin America: Geographic Structure, Phenotypic Diversity and Self-Perception of Ancestry Based on 7,342 Individuals". PLOS Genetics. 10 (9): e1004572. Bibcode:2014PLOSG..10.4572R. doi:10.1371/journal.pgen.1004572. ISSN 1553-7390. PMC 4177621. PMID 25254375.
Muiños Díaz, Yerena; Saornil, Maria A.; Almaraz, Ana; Muñoz-Moreno, Mari F.; García, Ciro; Sanz, Ruperto (2009). "Iris color: validation of a new classification and distribution in a Spanish population-based sample". European Journal of Ophthalmology. 19 (4): 686–689. doi:10.1177/112067210901900427. ISSN 1120-6721. PMID 19551689. S2CID 40940828.
Lucock, Mark D. (25 June 2022). "The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and <scp>UV</scp> radiation during human expansion". American Journal of Biological Anthropology. 180 (2). Wiley: 252–271. doi:10.1002/ajpa.24564. ISSN 2692-7691. PMC 10083917. PMID 36790744. This may also be a direct counter‐measure to a short winter photoperiod at high latitudes; blue eyes increase intraocular light scattering and thereby suppress melatonin release from the pineal gland. This may be an adaptive trait to reduce/prevent depression, a condition linked to short day length (Higuchi et al., 2007; Lucock et al., 2021).
Belkin, Douglas (18 October 2006). "Blue eyes are increasingly rare in America - Americas - International Herald Tribune (Published 2006)". The New York Times. ISSN 0362-4331. Retrieved 2 February 2021.
O'Connor, Anahad (19 January 2009). "The Claim: Eye Color Can Have an Effect on Vision". The New York Times. ISSN 0362-4331. Retrieved 5 May 2021.
Miller, L. K.; Rowe, P. J.; Lund, J. (August 1992). "Correlation of eye color on self-paced and reactive motor performance". Perceptual and Motor Skills. 75 (1): 91–95. doi:10.2466/pms.1992.75.1.91. ISSN 0031-5125. PMID 1528697. S2CID 31398375.
Rowe, P. J.; Evans, P. (August 1994). "Ball color, eye color, and a reactive motor skill". Perceptual and Motor Skills. 79 (1 Pt 2): 671–674. doi:10.2466/pms.1994.79.1.671. ISSN 0031-5125. PMID 7808908. S2CID 6743916.
Crowe, M.; O'Connor, D. (October 2001). "Eye colour and reaction time to visual stimuli in rugby league players". Perceptual and Motor Skills. 93 (2): 455–460. doi:10.2466/pms.2001.93.2.455. ISSN 0031-5125. PMID 11769902. S2CID 22785194.