Eye color - Knowledge

501: 573: 2403:"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. 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 (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." 525:(also found in green eyes). Amber eyes should not be confused with hazel eyes. Although hazel eyes may contain specks of amber or gold, they usually tend to have many other colors, including green, brown and orange. Also, hazel eyes may appear to shift in color and consist of flecks and ripples, while amber eyes are of a solid gold hue. Even though amber is similar to gold, some people have russet or copper colored amber eyes that are mistaken for hazel, though hazel tends to be duller and contains green with red/gold flecks, as mentioned above. Amber eyes may also contain amounts of very light gold-ish gray. 740: 1473: 1502:. However, there is a study that found that dark-eyed people perform better at "reactive-type tasks", which suggests they may have better reaction times. People with light-colored eyes, however, performed better at so-called "self-paced tasks", which include activities like hitting a golf ball or throwing baseballs. In another study, people with darker eyes performed better at hitting racquetballs. There are also other studies that challenge these findings. According to scientists, more study is needed to verify these results. 990: 1037:(which is not strongly frequency-dependent) rather than Rayleigh scattering (in which shorter wavelengths of light are scattered more). This would be analogous to the change in the color of the sky, from the blue given by the Rayleigh scattering of sunlight by small gas molecules when the sky is clear, to the gray caused by Mie scattering of large water droplets when the sky is cloudy. Alternatively, it has been suggested that gray and blue eyes might differ in the concentration of melanin at the front of the stroma. 1021: 410: 974: 40: 1088: 1303: 1266:, the color of the iris is typically blue but can vary from blue to brown. In severe forms of albinism, there is no pigment on the back of the iris, and light from inside the eye can pass through the iris to the front. In these cases, the only color seen is the red from the hemoglobin of the blood in the capillaries of the iris. Such albinos have pink eyes, as do albino rabbits, mice, or any other animal with a total lack of melanin. 1364: 940: 637: 398: 1356: 581: 960: 775: 764: 514: 732: 1111: 548:, also known as lipochrome. Many animals such as canines, domestic cats, owls, eagles, pigeons and fish have amber eyes, whereas in humans this color occurs less frequently. Amber is the third-rarest natural eye color after green and gray, in 5% of the world's population. People with that eye color are found in 1179: 1007:

in the United States to be 33.8% for those born from 1936 through 1951, compared with 57.4% for those born from 1899 through 1905. As of 2006, one out of every six Americans, or 16.6% of the total US population, has blue eyes, including 22.3% of whites. The incidence of blue eyes continues to decline

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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

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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.;

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and a moderate amount of melanin in the iris' anterior border layer. Hazel eyes often appear to shift in color from a brown to a green. Although hazel mostly consists of brown and green, the dominant color in the eye can either be brown/gold or green. This is why hazel eyes can be mistaken as amber,

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Almost all mammals have brown or darkly-pigmented irises. In humans, brown is by far the most common eye color, with approximately 79% of people in the world having it. Brown eyes result from a relatively high concentration of melanin in the stroma of the iris, which causes light of both shorter and

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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

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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.

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Normal eye colors range from the darkest shades of brown to the lightest tints of blue. To meet the need for standardized classification, at once simple yet detailed enough for research purposes, Seddon et al. developed a graded system based on the predominant iris color and the amount of brown or

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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".

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Iris color can provide a large amount of information about a person, and a classification of colors may be useful in documenting pathological changes or determining how a person may respond to ocular pharmaceuticals. Classification systems have ranged from a basic light or dark description to

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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

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Normally, there is a thick layer of melanin on the back of the iris. Even people with the lightest blue eyes, with no melanin on the front of the iris at all, have dark brown coloration on the back of it, to prevent light from scattering around inside the eye. In those with milder forms of

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The green color is caused by the combination of: 1) an amber or light brown pigmentation in the stroma of the iris (which has a low or moderate concentration of melanin) with: 2) a blue shade created by the Rayleigh scattering of reflected light. Green eyes contain the yellowish pigment

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was born with one blue eye and one green eye, a trait that reportedly was common in his family, suggesting that it was a genetic trait. This anomaly, which film producers thought would be disturbing to film audiences, was "corrected" by having Van Cleef wear brown contact lenses.

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and why amber is often counted as hazel in studies, and vice versa. The combination can sometimes produce a multicolored iris, i.e., an eye that is light brown/amber near the pupil and charcoal or dark green on the outer part of the iris (or vice versa) when observed in sunlight.

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The most important role of melanin in the iris is to protect the eyes from the sun's harmful rays. People with lighter eye colors, such as blue or green, have lessened protection from the sun, and so need greater protection from the sun's rays than those with darker eye colors.

794:. Unlike brown eyes, blue eyes have low concentrations of melanin in the stroma of the iris, which lies in front of the dark epithelium. Longer wavelengths of light tend to be absorbed by the dark underlying epithelium, while shorter wavelengths are reflected and undergo 1381:) is an eye condition in which one iris is a different color from the other (complete heterochromia), or where a part of one iris is a different color from the remainder (partial heterochromia or sectoral heterochromia). It is a result of the relative excess or lack of 2233:"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 , Holland , Australia or Poland , as well as in this study (see Fig. 2)." 845:. The same DNA sequence of the OCA2 gene among blue-eyed people suggests they may have a single common ancestor. The researchers hypothesized that the OCA2 mutation responsible for blue eyes arose in an individual who lived in the northwestern part of the 1497:

Although people with lighter eye color are generally more sensitive to light because they have less pigment in the iris to protect them from sunlight, there is little to no evidence that eye color has a direct impact on vision qualities such as

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Like blue eyes, gray eyes have a dark epithelium at the back of the iris and a relatively clear stroma at the front. One possible explanation for the difference in the appearance of gray and blue eyes is that gray eyes have larger deposits of

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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".

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In humans, eye color is a highly sexually dimorphic trait. Several studies have shown that men are more likely to have blue eyes than women, while women are more likely to have darker eye colors (green and brown eyes) than men.

293:. A 2010 study of eye color variation in hue and saturation values using high-resolution digital full-eye photographs found three new loci for a total of ten genes, allowing the explanation of about 50% of eye color variation. 2188:

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".

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can appear purple or violet at certain times, "true" violet-colored eyes occur only due to albinism. Eyes that appear red or violet under certain conditions due to albinism are less than 1 percent of the world's population.

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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".

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due to lack of iridial pigmentation. The ocular albino also lacks normal amounts of melanin in the retina as well, which allows more light than normal to reflect off the retina and out of the eye. Because of this, the

96:(located on the back of the iris), the melanin content within the iris stroma (located at the front of the iris), and the cellular density of the stroma. The appearance of blue, green, and hazel eyes results from the 2278:

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).

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The limbal ring is also a feature of the iris contributing to eye color, appearing as a darkened, occasionally black region encircling the iris resulting from a manifestation of the optical properties of the

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research has identified human remains much older than the Neolithic period with the OCA2 mutation for blue eyes. It is now believed that the OCA2 allele dates back to the recent migration of modern humans

2268:"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)." 1538:

in the first half of the 20th century. The scale consists of 20 colors ranging from light blue to dark brown-black, corresponding to natural eye colors caused by the amount of melanin in the iris:

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17% of children experienced a change of eye color from early childhood to adulthood, and of those children, 50% of developed lighter eyes as they got older. The other 50% developed darker eyes.

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Generally, children with hazel and light brown eyes tended to experience a lightening of their eye color by adulthood. Children with green eyes often experienced a darkening of their eye color.

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Around 18% of the US population and 5% of the world population have hazel eyes. 55.2% of Spanish subjects in a series of 221 photographs were judged to have hazel eyes. Hazel eyes are found in

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The genetics and inheritance of eye color in humans is complicated. As of 2010, as many as 16 genes have been associated with eye color inheritance. Some of the eye-color genes include

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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.

1102:, it is also possible to have two different eye colors. This occurs in humans and certain breeds of domesticated animals and affects less than 1 percent of the world's population. 2722: 4157: 1546:. Green irises, for example, have some yellow and the blue structural color. Brown irises contain more or less melanin. Some eyes have a dark ring around the iris, called a 898:

It has been proposed that blue eyes may have been adaptive to shorter day lengths at higher latitudes, as blue eyes increase inferocular light scattering, which suppresses

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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".

3670:"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." 215:

There is evidence that as many as 16 different genes could be responsible for eye color in humans; however, the main two genes associated with eye color variation are

1158:(ARMD) than those with darker iris color; lighter eye color is also associated with an increased risk of ARMD progression. A gray iris may indicate the presence of a 1969:

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).

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depends on viewing conditions (e.g., the amount and kind of illumination, as well as the hue of the surrounding environment), so does the perception of eye color.

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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".

5054: 2971:; Gillespie, Nathan A.; Ewen, Kelly R.; Jewell, Mary; Liew, Yew Wah; Hayward, Nicholas K.; Sturma, Richard A.; Trenta, Jeffrey M.; Martina, Nicholas G. (2004). 1443:

affecting the development of one eye, possibly through some sort of genetic mutation. Occasionally, heterochromia can be a sign of a serious medical condition.

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experienced an eye color change during the same period, with most developing lighter eyes, relative to their original color at the time of their child's birth.

3757:"The evolution of human skin pigmentation: A changing medley of vitamins, genetic variability, and <scp>UV</scp> radiation during human expansion" 2022: 1166:

has been found in those with blue, green or gray eyes. However, a study in 2000 suggests that people with dark brown eyes are at increased risk of developing

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Kayser, Manfred; Liu, Fan; Janssens, A. Cecile J.W.; Rivadeneira, Fernando; Lao, Oscar; Van Duijn, Kate; Vermeulen, Mark; Arp, Pascal; et al. (2008).

1134:", in which the very bright light from a flash reflects off the retina, which is abundantly vascular, causing the pupil to appear red in the photograph. 150:

has been shown to be incorrect. The genetics of eye color are so complex that almost any parent-child combination of eye colors can occur. However, OCA2

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Rafnsson V, Hrafnkelsson J, Tulinius H, Sigurgeirsson B, Olafsson JH (2004). "Risk factors for malignant melanoma in an Icelandic population sample".

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yellow pigment present. There are three pigment colors that determine, depending on their proportion, the outward appearance of the iris, along with

1485:. Limbal rings are not present in all individuals, and their thickness and prominence may correlate with health or youthfulness, and contributes to 5126: 1458:, can also cause increased pigmentation in one eye. On occasion, a difference in eye color is caused by blood staining the iris after injury. 1012:, 56% have blue/green eyes. In a series of 221 photographs of Spanish subjects, 16.3% of the subjects were determined to have blue-gray eyes. 3733: 1208:

in cells. Instead, the copper accumulates in the liver and in other tissues, including the iris of the eye. This results in the formation of

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detailed gradings employing photographic standards for comparison. Others have attempted to set objective standards of color comparison.

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Piquet-Thepot M.-M. - Bulletins et Mémoires de la Société d'anthropologie de Paris, XII° Série, tome 3 fascicule 3, pg. 207,208 - (1968)

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is therefore a major factor in the expression of eye color genotypes. One study suggested that women's higher levels of the sex hormone

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Hawkins TA, Stewart WC, McMillan TA, Gwynn DR (1994). "Analysis of diode, argon, and Nd: YAG peripheral iridectomy in cadaver eyes".

4106: 2281:"Sex-specific genetic effects associated with pigmentation, sensitivity to sunlight, and melanoma in a population of Spanish origin" 1003:

Approximately 8% to 10% of the global population have blue eyes. A 2002 study found that the prevalence of blue eye color among the

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technology have revealed some of the history of eye color in Europe. Through the analysis of ancient DNA, a 2020 study published in

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Amber eyes are a solid color with a strong yellowish/golden and russet/coppery tint, which may be due to the yellow pigment called

1436:, on the other hand, had the appearance of different eye colors due to an injury that caused one pupil to be permanently dilated. 973: 277:, where they may influence the expression of the gene product, which in turn affects pigmentation. A specific mutation within the 4400:

Stang A, Ahrens W, Anastassiou G, Jöckel KH (2003). "Phenotypical characteristics, lifestyle, social class and uveal melanoma".

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appears with gray eyes (γλαυκῶπις). Under magnification, gray eyes exhibit small amounts of yellow and brown color in the iris.

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Albert, Daniel M; Green, W Richard; Zimbric, Michele L; Lo, Cecilia; Gangnon, Ronald E; Hope, Kirsten L; Gleiser, Joel (2003).

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is a congenital condition characterized by an extremely underdeveloped iris, which appears absent on superficial examination.

544:(called xanthophores) located in the iris stroma. In humans, yellowish specks or patches are thought to be due to the pigment 177:

or SNPs. The total number of genes that contribute to eye color is unknown, but there are a few likely candidates. A study in

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Miller, L. K.; Rowe, P. J.; Lund, J. (August 1992). "Correlation of eye color on self-paced and reactive motor performance".

4580: 4547: 4212: 3829: 2689: 2254: 1773: 1721: 572: 3985: 5090: 4078: 181:(2009) found that it was possible to predict eye color with more than 90% accuracy for brown and blue using just six SNPs. 5189: 2742: 3862: 3845: 2704: 250:. (The name of the gene is derived from the disorder it causes, oculocutaneous albinism type II.) Different SNPs within 3542: 155: 5058: 3918: 1420:

can have two different colored eyes just like any two siblings can—because each cell has different eye color genes. A

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Sturm R.A. & Larsson M., Genetics of human iris colour and patterns, Pigment Cell Melanoma Res, 22:544-562, 2009.

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medium of the stroma. This is the same scattering that accounts for the blue appearance of the sky. The result is a "

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Crowe, M.; O'Connor, D. (October 2001). "Eye colour and reaction time to visual stimuli in rugby league players".

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People of European descent show the greatest variety in eye color of any population worldwide. Recent advances in

4322:; McCarty, Catherine A (2003). "Iris colour, ethnic origin and progression of age-related macular degeneration". 3305:

Muiños Díaz, Yerena; Saornil, Maria A.; Almaraz, Ana; Muñoz-Moreno, Mari F.; García, Ciro; Sanz, Ruperto (2009).

3623: 3561:"Is there any difference in the photobiological properties of melanins isolated from human blue and brown eyes?" 2073:

Grant MD, Lauderdale DS (2002). "Cohort effects in a genetically determined trait: eye colour among US whites".

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Oliphant LW (1981). "Crystalline pteridines in the stromal pigment cells of the iris of the great horned owl".

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Blue eyes with a brown spot, green eyes, and gray eyes are caused by an entirely different part of the genome.

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de la Maza, Maite Sainz; Tauber, Joseph; Foster, C. Stephen (2012). "Noninflammatory Diseases of the Sclera".

1971:"A three-single-nucleotide polymorphism haplotype in intron 1 of OCA2 explains most human eye-color variation" 4134: 4286:"Melanins: Skin Pigments and Much More—Types, Structural Models, Biological Functions, and Formation Routes" 173:. These genes are sought by studying small changes in the genes themselves and in neighboring genes, called 4771:"Put a (Limbal) Ring on It: Women Perceive Men's Limbal Rings as a Health Cue in Short-Term Mating Domains" 3440:

variation) showed that at least 60% of the ancient Siberian specimens under study had blue (or green) eyes.

1941:"Eye color definition – Medical Dictionary definitions of popular medical terms easily defined on MedTerms" 1666:"Characterization of melanins in human irides and cultured uveal melanocytes from eyes of different colors" 906:, perhaps reducing psychological depression (which is linked to the short day length of higher latitudes). 4020:, Volumes 16–17 By Société de statistique, d'histoire et d'archéologie de Marseille et de Provence p. 273 4563:

Wheatley, TJ (2006). "Upper gastrointestinal surgery". In Kingsnorth, Andrew N; Majid, Aljafri A (eds.).

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There are many other possible reasons for having two different-colored eyes. For example, the film actor

391: 2415:"Three genome-wide association studies and a linkage analysis identify HERC2 as a human iris color gene" 3877:"Blue eyes are increasingly rare in America - Americas - International Herald Tribune (Published 2006)" 2183: 2181: 2179: 2177: 1320: 757: 4716: 3846:"Distribution of Bodily Characters. Pigmentation, the Pilous System, and Morphology of the Soft Parts" 3307:"Iris color: validation of a new classification and distribution in a Spanish population-based sample" 1074:

Gray is the second-rarest natural eye color after green, with 3% of the world's population having it.

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located a single mutation that causes the phenomenon of blue eyes. The research was published in the

17: 2973:"A genome scan for eye color in 502 twin families: most variation is due to a QTL on chromosome 15q" 2174: 1553:

Eye color in non-human animals is regulated differently. For example, instead of blue as in humans,

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expression, is partly responsible for blue eyes. Other genes implicated in eye color variation are

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The brightly colored eyes of many bird species result from the presence of other pigments, such as

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

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which accounts for the blue sky. Neither blue nor green pigments are present in the human iris or

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vary: it is sometimes considered to be synonymous with light brown or gold, as in the color of a

93: 5218: 3097:"Induction of tyrosinase gene transcription in human iris organ cultures exposed to latanoprost" 4443:

Cumming RG, Mitchell P, Lim R (2000). "Iris color and cataract: The Blue Mountains Eye Study".

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In many parts of the world, it is nearly the only iris color present. Brown eyes are common in

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Ask a Geneticist: If both parents have blue eyes, how could they have a child with brown eyes?

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can have two different colored eyes if the DNA difference happens to be in an eye-color gene.

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found that eye color may be subject to change in infancy, and from adolescence to adulthood.

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Nicolas, Caroline M; Robman, Luba D; Tikellis, Gabriella; Dimitrov, Peter N; Dowrick, Adam;

3818: 2679: 2518:"Digital Quantification of Human Eye Color Highlights Genetic Association of Three New Loci" 2134:

Liu, Fan; Van Duijn, Kate; Vingerling, Johannes R.; Hofman, Albert; Uitterlinden, André G.;

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Wielgus AR, Sarna T (2005). "Melanin in human irides of different color and age of donors".

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Rowe, P. J.; Evans, P. (August 1994). "Ball color, eye color, and a reactive motor skill".

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Hammond BR, Fuld K, Snodderly DM (1996). "Iris color and macular pigment optical density".

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Cavalli-Sforza, Luigi Luca; Cavalli-Sforza, Luca; Menozzi, Paolo; Piazza, Alberto (1994).

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may appear red under certain lighting conditions owing to the extremely low quantities of

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An example of sectoral heterochromia. The subject has a blue iris with a brown section.

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An example of complete heterochromia. The subject has one brown eye and one hazel eye.

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Percentage of dark eyes among British military recruits in the 19th century by region

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Another hypothesis about heterochromia is that it can result from a viral infection

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Hejkal TW, Camras CB (1999). "Prostaglandin analogs in the treatment of glaucoma".

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to establish the eye color of an individual. It was created by the anthropologists

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Eye color outside of the iris may also be symptomatic of disease. Yellowing of the

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Oliphant LW (1987). "Pteridines and purines as major pigments of the avian iris".

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Blue eyes are predominant in northern and eastern Europe, particularly around the

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in the stroma, so that the light that is reflected from the epithelium undergoes

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have shown that men are substantially more likely to have blue eyes than women.

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Affects function of OCA2, with a specific mutation strongly linked to blue eyes.

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Green eyes probably result from the interaction of multiple variants within the

5248: 5186: 4736: 3911:"Don't it make my blue eyes brown Americans are seeing a dramatic color change" 3322: 3201: 3184: 3113: 3096: 2989: 2972: 2739: 2430: 1743: 1482: 1447: 1280: 1163: 1131: 1034: 959: 803: 713: 697: 693: 616: 486: 426: 361: 97: 85: 78: 66: 5120:

Seddon, JM; CR Sahagian; RJ Glynn; RD Sperduto; ES Gragoudas (1 August 1990).

4965: 4914: 4693: 3849: 3694: 3418: 2903: 2297: 2158: 2139: 2086: 2041: 5227: 5091:"Contribution à l'anthropologie des Corses: Anthropologie de la tête (suite)" 4973: 4922: 4871: 4863: 4826: 4787: 4770: 4613: 3934:"Prediction of eye color in the Slovenian population using the IrisPlex SNPs" 3888: 3780: 3330: 2968: 2932: 2857: 2382: 2316: 2220: 2120: 1642: 1499: 1428: 1154:

Those with lighter iris color have been found to have a higher prevalence of

1044: 824:

The inheritance pattern followed by blue eyes was previously assumed to be a

783: 561: 541: 228: 5106: 4717:"Preliminary evidence that the limbal ring influences facial attractiveness" 4495: 3949: 3539: 3306: 2841: 2631: 2277: 2246:

Forensic DNA Analysis: Technological Development and Innovative Applications

1871: 5238: 4981: 4796: 4755: 4701: 4513: 4464: 4421: 4386: 4343: 4319: 4262: 3967: 3932:

Kastelic, V; Pośpiech, E; Draus-Barini, J; Branicki, W; Drobnič, K (2013).

3798: 3702: 3511: 3426: 3338: 3280: 3210: 3166: 3122: 3081: 3039: 2998: 2950: 2759:

Oliphant LW (1987). "Observations on the pigmentation of the pigeon iris".

2632:

Blue eyes in lemurs and humans: same phenotype, different genetic mechanism

2553: 2493: 2448: 2390: 2334: 2228: 2167: 2094: 2049: 2004: 1914: 1682: 1665: 1650: 1574: 1519: 1060: 1020: 926: 918: 903: 557: 537: 490: 477:. Light brown eyes bordering amber and hazel coloration are more common in 462: 430: 409: 204:

suggested that the common gene for blue eye color likely originated in the

31: 5139: 5027: 4930: 4879: 4664: 4651: 4302: 4285: 3594: 3470: 3377:

The Races of Britain. A contribution to the anthropology of Western Europe

3288: 3245: 3057: 3014:"Iris melanocyte numbers in Asian, African American, and Caucasian irides" 2815: 2780: 2588: 1857: 1765: 1691: 1279:

is much more pronounced in albino individuals, and this can emphasize the

1170:

and therefore should protect their eyes from direct exposure to sunlight.

254:

are strongly associated with blue and green eyes as well as variations in

39: 4364: 4253: 4236: 3815: 3576: 1905: 1888: 1748:

Tenth IEEE International Conference on Computer Vision (ICCV'05) Volume 1

1741: 1601: 1547: 1523: 1467: 1451: 1433: 1052: 965: 858: 768: 705: 612: 553: 494: 474: 446: 197: 3559:

Menon IA, Basu PK, Persad S, Avaria M, Felix CC, Kalyanaraman B (1987).

1822: 1401:

content. A number of causes are responsible, including genetic, such as

1178: 3772: 3503: 3237: 2807: 2485: 2307: 2211: 1586: 1327: in this section. Unsourced material may be challenged and removed. 1087: 1064: 922: 910: 787: 725: 682: 545: 522: 470: 434: 341: 290: 263: 128: 124: 70: 3488:"Genetic determinants of hair, eye and skin pigmentation in Europeans" 2516:

Martin, Nicholas G.; Hammond, Christopher J.; Kayser, Manfred (2010).

2373: 2356: 1236:. A blue coloration of the sclera may also be symptomatic of disease. 832:, meaning that it is controlled by the interactions of several genes. 528:

The eyes of some pigeons contain yellow fluorescing pigments known as

4715:

Darren Peshek; Negar Semmaknejad; Donald Hoffman; Pete Foley (2011).

4618: 4601: 3631: 3304: 1596: 1557: 1233: 1229: 1056: 930: 899: 854: 846: 799: 620: 529: 482: 466: 453:. Light or medium-pigmented brown eyes can also be commonly found in 438: 422: 255: 205: 178: 116: 74: 5119: 3931: 1302: 88:

varies from light brown to black, depending on the concentration of

3627: 3058:"Iris color and intraocular pressure: the Blue Mountains Eye Study" 2140:"Eye color and the prediction of complex phenotypes from genotypes" 1986: 1554: 1363: 1263: 1250: 1245: 1225: 1167: 1030: 1009: 950: 601: 458: 450: 190: 51: 1450:, which can result in a number of heterochromatic traits, such as 5044:

Technical report 1495, University of Wisconsin–Madison, Dec 2003.

2967:

Zhu, Gu; Evans, David M.; Duffy, David L.; Montgomery, Grant W.;

1398: 1390: 1382: 1159: 1123: 1041: 791: 701: 678: 442: 374: 331: 313: 286: 89: 58: 4018:

Provincia: bulletin trimestriel de la Société de Statistique ...

3865:. Adelaidenow.com.au (11 January 2008). Retrieved on 2011-12-23. 2708: 1664:

Prota G, Hu DN, Vincensi MR, McCormick SA, Napolitano A (1998).

1355: 1212:, which are dark rings that encircle the periphery of the iris. 866:

roughly 50,000 years ago, and entered Europe from western Asia.

397: 239: 208:

and arrived in Europe around 42,000 years ago, after the exodus

2470: 1930:. Sciencedaily.com (22 February 2007). Retrieved on 2011-12-23. 1394: 1221: 1201: 1068: 946: 874: 850: 818: 608: 580: 549: 536:

are thought to be due to the presence of the pteridine pigment

478: 454: 418: 120: 3679: 3095:

Lindsey JD, Jones HL, Hewitt EG, Angert M, Weinreb RN (2001).

4317: 4033: 3403: 3056:

Mitchell R, Rochtchina E, Lee A, Wang JJ, Mitchell P (2003).

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trait, however, eye color inheritance is now recognized as a

774: 763: 513: 321: 223: 142: 5095:

Bulletins et Mémoires de la Société d'anthropologie de Paris

4399: 2412: 2133: 1887:

White, Désirée; Rabago-Smith, Montserrat (14 October 2010).

1110: 731: 3223: 2840:

Lefohn A, Budge B, Shirley P, Caruso R, Reinhard E (2003).

2709:

SKIN/HAIR/EYE PIGMENTATION, VARIATION IN, 1; SHEP1 - 227220

2567:

Bito, LZ (1997). "Eye Color Changes Past Early Childhood".

1126:, allowing the blood vessels to show through. In addition, 782:

There is no blue pigmentation either in the iris or in the

673: 307: 217: 170: 136: 43:

The irises of human eyes exhibit a wide spectrum of colours

3809: 3549:. allaboutgenes.weebly.com. Retrieved on 8 September 2016. 2514: 2023:"Eye colour: portals into pigmentation genes and ancestry" 619:, and have also been observed to be very common among the 193:

may explain why women tend to have darker eyes than men.

30:"Iris color" redirects here. For the shade of purple, see 5159: 3055: 346:

Associated with differences between blue and green eyes.

336:

Associated with differences between blue and green eyes.

242:), when in a variant form, causes the pink eye color and 186: 62: 5005: 3141:"Race, iris color, and age-related macular degeneration" 3094: 2839: 1663: 1397:. This uncommon condition usually results due to uneven 869:

Eiberg and colleagues suggested in a study published in

3607: 3182: 2966: 2612: 1968: 853:

sometime between 6,000 and 10,000 years ago during the

169:

Eye color is an inherited trait determined by multiple

4529: 3986:"Are gray eyes the same as blue in terms of genetics?" 3185:"Iris color as a prognostic factor in ocular melanoma" 3138: 2602: 2600: 2598: 1196:

involves a mutation of the gene coding for the enzyme

4237:"Genotype–phenotype associations and human eye color" 3558: 3018:

Transactions of the American Ophthalmological Society

2749:. Eyecarecontacts.com. Retrieved on 23 December 2011. 1889:"Genotype–phenotype associations and human eye color" 677:

and other genes. They may have been present in south

146:. The earlier belief that blue eye color is a simple 5196:. Edromanguitars.com. Retrieved on 23 December 2011. 3461:. Eyedoctorguide.com. Retrieved on 23 December 2011. 3266: 3011: 5122:"Evaluation of an iris color classification system" 4815:"The Claim: Eye Color Can Have an Effect on Vision" 4199:(Second ed.). Academic Press. pp. 24–26. 3990:

Understanding Genetics: Human Health and the Genome

2630:Brenda J Bradley, Anja Pedersen, Nicholas I Mundy: 2595: 2355:Hanel, Andrea; Carlberg, Carsten (September 2020). 1228:, and may be symptomatic of liver diseases such as 1137:Although the deep blue eyes of some people such as 712:, green eyes are most common among those of recent 4629: 4442: 3817: 3654:"How one ancestor helped turn our brown eyes blue" 3471:Why Do Europeans Have So Many Hair and Eye Colors? 3452:Blue Eyes Versus Brown Eyes: A Primer on Eye Color 316:producing cells. Central importance to eye color. 4951: 4768: 4235:White, Désirée; Rabago-Smith, Montserrat (2010). 4234: 3486:Sulem, P.; Gudbjartsson, D.; et al. (2007). 3485: 3473:. Cogweb.ucla.edu. Retrieved on 23 December 2011. 3183:Regan S, Judge HE, Gragoudas ES, Egan KM (1999). 2842:"An Ocularist's Approach to Human Iris Synthesis" 1886: 895:with subsequent reduction in melanin production. 883:gene, which is hypothesized to interact with the 390:"Brown eyes" redirects here. For other uses, see 5225: 5127:Investigative Ophthalmology & Visual Science 4477: 3726:"Blue eyes result of ancient genetic 'mutation'" 3139:Frank RN, Puklin JE, Stock C, Canter LA (2000). 2720:Howard Hughes Medical Institute: Ask A Scientist 2116:"DNA test for eye colour could help fight crime" 1872:"Phenotypes and Genotypes for human eye colors." 1446:A common cause in females with heterochromia is 1385:within an iris or part of an iris, which may be 1270:defects can almost always be observed during an 1256: 756:"Blue eyes" redirects here. For other uses, see 100:of light in the stroma, a phenomenon similar to 4155: 3717: 2072: 1943:. Medterms.com. 29 October 2003. Archived from 1928:No Single Gene For Eye Color, Researchers Prove 1454:. Trauma and certain medications, such as some 817:eye color. Studies from various populations in 810:that varies with external lighting conditions. 4849: 4149: 4113:. Albinism.org. Retrieved on 23 December 2011. 3624:"Genetic mutation makes those brown eyes blue" 2681:Biochromy: Natural Coloration of Living Things 1882: 1880: 1713:Biochromy: Natural Coloration of Living Things 1224:(the "whites of the eyes") is associated with 5162:"Subspecies comparison of the Genus: Corucia" 5088: 5042:Quantification and Correction of Iris Color." 4762: 3992:. Stanford School of Medicine. Archived from 3979: 3977: 3673: 2684:. University of California Press. p. 9. 2354: 2020: 1874:Athro Limited website. Retrieved 10 May 2006. 700:. Around 8–10% of men and 18–21% of women in 373:It was also found that 11% of the children's 4812: 4679: 4122: 3804:(Higuchi et al., 2007; Lucock et al., 2021). 1620: 4308:Eye color change and the affecting factors. 3983: 3902: 3838: 3761:American Journal of Biological Anthropology 3051: 3049: 2793: 2758: 1877: 1835: 1215: 161:, explains most human eye-color variation. 4186: 3974: 2729:. Hhmi.org. Retrieved on 23 December 2011. 2242: 1505: 623:speaking populations of northern Germany. 5001: 4999: 4900: 4786: 4745: 4735: 4525: 4523: 4503: 4471: 4301: 4252: 3957: 3788: 3723: 3584: 3397: 3380:. Michigan State University. p. 39. 3300: 3298: 3200: 3156: 3112: 3029: 2988: 2940: 2922: 2835: 2833: 2543: 2533: 2438: 2372: 2324: 2306: 2296: 2210: 2157: 1994: 1904: 1829: 1755: 1705: 1703: 1701: 1681: 1343:Learn how and when to remove this message 998:, also known as the blue-eyed black lemur 790:is brownish black due to the presence of 4562: 4358: 4180: 4097: 3820:The History and Geography of Human Genes 3617: 3615: 3481: 3479: 3392:the Saxons...being very often hazel-eyed 3178: 3176: 3134: 3132: 3046: 2466: 2464: 2462: 2460: 2458: 2350: 2348: 2191:Forensic Science International: Genetics 2068: 2066: 2016: 2014: 1476:Light brown iris with a dark limbal ring 1471: 1362: 1354: 1177: 1118:The eyes of people with severe forms of 1109: 1086: 1019: 835:In 2008, a team of researchers from the 773: 762: 738: 730: 579: 571: 512: 499: 408: 396: 164: 38: 5113: 4436: 4311: 4187:Fertl, Dagmar; Rosel, Patricia (2009). 3260: 1964: 1962: 1804: 1802: 1145: 1082: 588:Hazel eyes are due to a combination of 355: 14: 5226: 5089:Piquet-Thepot, M.-M. (26 March 1968). 4996: 4623: 4520: 4283: 3908: 3874: 3754: 3621: 3373: 3295: 2962: 2960: 2830: 2673: 2671: 2406: 2357:"Skin colour and vitamin D: An update" 1737: 1735: 1733: 1698: 1614: 1091:Two different eye colors are known as 1040:Gray eyes can also be found among the 704:and 6% of men and 17% of women in the 4845: 4843: 4808: 4806: 4393: 4079:"The World's Population By Eye Color" 4073: 4071: 4022:l'iris gris est celui des chaouias... 3612: 3552: 3476: 3173: 3129: 2648: 2644: 2642: 2640: 2455: 2345: 2249:. Apple Academic Press. p. 207. 2063: 2011: 1864: 1809:Mason, Clyde W. (1924). "Blue Eyes". 1808: 1742:Huiqiong Wang; Lin, S.; Xiaopei Liu; 127:. Humans and other animals have many 5160:Jones, S.L.; Schnirel, B.L. (2006). 4813:O'Connor, Anahad (19 January 2009). 4158:"Elizabeth Taylor: Beautiful Mutant" 3748: 3724:Highfield, Roger (30 January 2008). 2618: 2606: 2581:10.1001/archopht.1997.01100150661017 2566: 2271: 2236: 1959: 1799: 1657: 1325:adding citations to reliable sources 1296: 630: 69:and the frequency-dependence of the 5034: 4708: 3909:Belkin, Douglas (17 October 2006). 3875:Belkin, Douglas (18 October 2006). 3361:"Where are hazel eyes most common?" 2957: 2705:Online Mendelian Inheritance in Man 2677: 2668: 1730: 1709: 1492: 1204:within the liver from entering the 1173: 413:A light brown iris with limbal ring 406:longer wavelengths to be absorbed. 84:In humans, the pigmentation of the 27:Polygenic phenotypic characteristic 24: 4840: 4803: 4205:10.1016/b978-0-12-373553-9.00006-7 4125:"HOW TO: Avoid the red eye effect" 4068: 3622:Bryner, Jeanna (31 January 2008). 2773:10.1111/j.1600-0749.1987.tb00414.x 2651:"Which Eye Colors Are the Rarest?" 2637: 1850:10.1111/j.1600-0749.1987.tb00401.x 1716:. University of California Press. 380: 25: 5260: 5203: 4614:Ocular Manifestations of Albinism 4565:Fundamentals of Surgical Practice 4445:American Journal of Ophthalmology 4156:Palmer, Roxanne (25 March 2005). 3311:European Journal of Ophthalmology 1105: 270:. The polymorphisms may be in an 5180: 5153: 5082: 4336:10.1046/j.1442-9071.2003.00711.x 4123:Dave Johnson (16 January 2009). 3755:Lucock, Mark D. (25 June 2022). 1635:10.1111/j.1600-0749.2005.00268.x 1301: 1286: 1156:age-related macular degeneration 1077: 988: 972: 958: 938: 635: 532:. The bright yellow eyes of the 5073: 5055:"Martin-Schulz Eye Color Chart" 5047: 4945: 4894: 4673: 4658: 4606: 4589: 4556: 4478:McDonnell G, Esmonde T (1999). 4277: 4228: 4116: 4043: 4026: 4008: 3925: 3868: 3856: 3646: 3601: 3533: 3464: 3445: 3367: 3353: 3217: 3088: 3005: 2897: 2872: 2787: 2752: 2732: 2713: 2698: 2624: 2560: 2508: 2127: 2109: 1933: 1921: 1560:eye color in the skink species 1312:needs additional citations for 175:single-nucleotide polymorphisms 57:determined by two factors: the 4197:Encyclopedia of Marine Mammals 3824:. Princeton University Press. 2021:Sturm RA, Frudakis TN (2004). 1790: 1461: 913:. Blue eyes are also found in 688:Green eyes are most common in 552:, and in fewer numbers in the 481:, but can also be observed in 13: 1: 5020:10.1016/S0275-5408(97)00070-7 4644:10.1016/S0039-6257(97)80018-5 4457:10.1016/S0002-9394(00)00479-7 3074:10.1016/S0002-9394(02)01967-0 2678:Fox, Denis Llewellyn (1979). 2243:Pilli, E.; Berti, A. (2021). 1811:Journal of Physical Chemistry 1750:. pp. 1691–1698 Vol. 2. 1710:Fox, Denis Llewellyn (1979). 1607: 1257:Ocular albinism and eye color 983:known to be born in captivity 857:period. However, more recent 596:Definitions of the eye color 5219:Eye Color and Human Diseases 4573:10.1017/CBO9780511545740.013 4324:Clin. Experiment. Ophthalmol 3151:: 109–15, discussion 115–7. 2924:10.1371/journal.pgen.1004572 2535:10.1371/journal.pgen.1000934 2203:10.1016/j.fsigen.2013.03.007 1566:is black, and the autosomal 1051:in Northwest Africa, in the 1008:among American children. Of 873:that a mutation in the 86th 281:gene, a gene that regulates 227:, and both are localized in 7: 5040:Fan S, Dyer CR, Hubbard L. 4954:Perceptual and Motor Skills 4903:Perceptual and Motor Skills 4852:Perceptual and Motor Skills 4665:Biography for Lee Van Cleef 4540:10.1007/978-1-4419-6502-8_8 4414:10.1076/opep.10.5.293.17319 4379:10.1016/j.ypmed.2004.03.027 2197:(4). Elsevier BV: 453–460. 1580: 1239: 1162:, and an increased risk of 646:may have misleading content 392:Brown eyes (disambiguation) 10: 5265: 4769:Brown M, Sacco DF (2018). 4737:10.1177/147470491100900201 3540:"OCA2: The Gene for Color" 3323:10.1177/112067210901900427 3202:10.1001/archopht.117.6.811 3114:10.1001/archopht.119.6.853 2990:10.1375/136905204323016186 2431:10.1016/j.ajhg.2007.10.003 2285:Biology of Sex Differences 2138:; Kayser, Manfred (2009). 1465: 1290: 1243: 758:Blue eyes (disambiguation) 755: 720:ancestry, with about 16%. 389: 29: 5172:(1): 1–25. Archived from 4966:10.2466/pms.2001.93.2.455 4915:10.2466/pms.1994.79.1.671 4694:10.3109/08820539909061464 4682:Seminars in Ophthalmology 4241:Journal of Human Genetics 3695:10.1007/s00439-007-0460-x 3419:10.1007/s00439-009-0683-0 2634:Am J Phys Anthropol. 2009 2569:Archives of Ophthalmology 2298:10.1186/s13293-016-0070-1 2159:10.1016/j.cub.2009.01.027 2087:10.1080/03014460210157394 2042:10.1016/j.tig.2004.06.010 1893:Journal of Human Genetics 1114:Red-appearing albino eyes 842:Journal of Human Genetics 131:variations in eye color. 4864:10.2466/pms.1992.75.1.91 4788:10.1177/0146167217733072 4104:NOAH – What is Albinism? 3732:. London. Archived from 2905:C. (25 September 2014). 2858:10.1109/MCG.2003.1242384 2846:IEEE Comput. Graph. Appl 2725:1 September 2010 at the 2361:Experimental Dermatology 2136:Janssens, A. Cecile J.W. 1216:Coloration of the sclera 891:, reduced expression of 837:University of Copenhagen 626: 567: 508: 385: 202:Experimental Dermatology 108:. This is an example of 5192:20 October 2006 at the 5107:10.3406/bmsap.1968.1417 4724:Evolutionary Psychology 4496:10.1136/pgmj.75.884.375 3950:10.3325/cmj.2013.54.381 3457:17 October 2008 at the 3145:Trans Am Ophthalmol Soc 2745:23 October 2010 at the 1506:Classification of color 1456:prostaglandin analogues 1130:can sometimes cause a " 1015: 813:Blue eyes are a highly 767:A light blue iris with 751: 708:have green eyes. Among 611:, most commonly in the 94:iris pigment epithelium 5008:Ophthalmic Physiol Opt 4290:New Journal of Science 3545:6 October 2016 at the 3281:10.1006/exer.1996.0035 1683:10.1006/exer.1998.0518 1570:color is yellow-green. 1477: 1368: 1360: 1210:Kayser–Fleischer rings 1190: 1115: 1095: 1025: 779: 771: 748: 736: 585: 577: 518: 505: 414: 402: 44: 4775:Pers Soc Psychol Bull 4296:. hindawi.com: 1–28. 3767:(2). Wiley: 252–271. 3374:Beddoe, John (1971). 1766:10.1109/ICCV.2005.215 1528:physical anthropology 1518:, developed from the 1487:facial attractiveness 1475: 1366: 1358: 1184:Kayser–Fleischer ring 1181: 1113: 1090: 1023: 777: 766: 742: 734: 583: 575: 516: 503: 412: 400: 165:Genetic determination 42: 5214:The Tech Interactive 4638:(Suppl 2): S117–23. 4567:. pp. 230–248. 4534:. pp. 277–297. 4480:"A homesick student" 4402:Ophthalmic Epidemiol 4254:10.1038/jhg.2010.126 4130:New Zealand PC World 3996:on 27 September 2011 3577:10.1136/bjo.71.7.549 2059:on 9 September 2006. 1906:10.1038/jhg.2010.126 1516:Martin–Schultz scale 1411:Waardenburg syndrome 1379:heterochromia iridis 1374:heterochromia iridum 1321:improve this article 1293:Heterochromia iridum 1146:Medical implications 1100:heterochromia iridum 1093:heterochromia iridum 1083:Two different colors 1067:. The Greek goddess 979:The first blue-eyed 356:Changes in eye color 302:Effect on eye color 154:, close to proximal 5176:on 2 February 2009. 4909:(1 Pt 2): 671–674. 4303:10.1155/2014/498276 4284:Solano, F. (2014). 4137:on 24 February 2010 4109:14 May 2012 at the 3921:on 22 October 2014. 3730:The Daily Telegraph 3363:. 26 February 2023. 1823:10.1021/j150239a007 1575:perception of color 826:mendelian recessive 796:Rayleigh scattering 786:. The iris pigment 747:with sea-green eyes 652:clarify the content 590:Rayleigh scattering 275:regulatory sequence 102:Rayleigh scattering 4819:The New York Times 3881:The New York Times 3773:10.1002/ajpa.24564 3736:on 1 November 2009 3504:10.1038/ng.2007.13 3238:10.1007/BF01203345 2808:10.1007/BF00233588 2621:, p. 661-662. 2486:10.1038/ng.2007.13 1522:, is one standard 1478: 1369: 1361: 1191: 1186:in a patient with 1116: 1096: 1026: 815:sexually dimorphic 780: 772: 749: 745:Chinchilla Persian 737: 710:European Americans 586: 578: 519: 506: 415: 403: 98:Tyndall scattering 79:stroma of the iris 45: 4582:978-0-511-54574-0 4549:978-1-4419-6501-1 4214:978-0-12-373553-9 3984:Lucy Southworth. 3831:978-0-691-08750-4 3660:. 31 January 2008 3608:Duffy et al. 2007 3498:(12): 1443–1452. 3062:Am. J. Ophthalmol 2969:Medland, Sarah E. 2691:978-0-520-03699-4 2649:Debrowski, Adam. 2419:Am. J. Hum. Genet 2374:10.1111/exd.14142 2256:978-1-000-06454-4 1975:Am. J. Hum. Genet 1775:978-0-7695-2334-7 1723:978-0-520-03699-4 1407:Horner's syndrome 1353: 1352: 1345: 1268:Transillumination 1200:, which prevents 1128:flash photography 949:with distinctive 902:release from the 669: 668: 350: 349: 159:regulatory region 152:gene polymorphism 16:(Redirected from 5256: 5197: 5187:Color Perception 5184: 5178: 5177: 5157: 5151: 5150: 5148: 5146: 5117: 5111: 5110: 5086: 5080: 5077: 5071: 5070: 5068: 5066: 5061:on 2 August 2016 5057:. Archived from 5051: 5045: 5038: 5032: 5031: 5003: 4994: 4993: 4949: 4943: 4942: 4898: 4892: 4891: 4847: 4838: 4837: 4835: 4833: 4810: 4801: 4800: 4790: 4766: 4760: 4759: 4749: 4739: 4721: 4712: 4706: 4705: 4677: 4671: 4662: 4656: 4655: 4627: 4621: 4610: 4604: 4593: 4587: 4586: 4560: 4554: 4553: 4527: 4518: 4517: 4507: 4475: 4469: 4468: 4440: 4434: 4433: 4397: 4391: 4390: 4362: 4356: 4355: 4315: 4309: 4307: 4305: 4281: 4275: 4274: 4256: 4232: 4226: 4225: 4223: 4221: 4184: 4178: 4177: 4175: 4173: 4168:on 27 March 2011 4164:. Archived from 4153: 4147: 4146: 4144: 4142: 4133:. Archived from 4120: 4114: 4101: 4095: 4094: 4092: 4090: 4085:. 6 October 2020 4075: 4066: 4065: 4063: 4061: 4055:All About Vision 4047: 4041: 4030: 4024: 4016: 4012: 4006: 4005: 4003: 4001: 3981: 3972: 3971: 3961: 3929: 3923: 3922: 3917:. Archived from 3915:The Boston Globe 3906: 3900: 3899: 3897: 3895: 3872: 3866: 3860: 3854: 3853: 3852:on 26 July 2011. 3848:. Archived from 3842: 3836: 3835: 3823: 3813: 3807: 3806: 3792: 3752: 3746: 3745: 3743: 3741: 3721: 3715: 3714: 3677: 3671: 3669: 3667: 3665: 3650: 3644: 3643: 3641: 3639: 3630:. Archived from 3619: 3610: 3605: 3599: 3598: 3588: 3556: 3550: 3537: 3531: 3530: 3528: 3526: 3483: 3474: 3468: 3462: 3449: 3443: 3442: 3401: 3395: 3394: 3371: 3365: 3364: 3357: 3351: 3350: 3302: 3293: 3292: 3264: 3258: 3257: 3221: 3215: 3214: 3204: 3189:Arch. Ophthalmol 3180: 3171: 3170: 3160: 3136: 3127: 3126: 3116: 3101:Arch. Ophthalmol 3092: 3086: 3085: 3053: 3044: 3043: 3033: 3009: 3003: 3002: 2992: 2964: 2955: 2954: 2944: 2926: 2901: 2895: 2894: 2892: 2890: 2884:All About Vision 2876: 2870: 2869: 2837: 2828: 2827: 2791: 2785: 2784: 2761:Pigment Cell Res 2756: 2750: 2736: 2730: 2717: 2711: 2702: 2696: 2695: 2675: 2666: 2665: 2663: 2661: 2655:All About Vision 2646: 2635: 2628: 2622: 2616: 2610: 2604: 2593: 2592: 2564: 2558: 2557: 2547: 2537: 2512: 2506: 2505: 2468: 2453: 2452: 2442: 2410: 2404: 2402: 2376: 2352: 2343: 2342: 2328: 2310: 2300: 2275: 2269: 2267: 2265: 2263: 2240: 2234: 2232: 2214: 2185: 2172: 2171: 2161: 2152:(5): R192–R193. 2131: 2125: 2113: 2107: 2106: 2070: 2061: 2060: 2058: 2052:. Archived from 2027: 2018: 2009: 2008: 1998: 1966: 1957: 1956: 1954: 1952: 1937: 1931: 1925: 1919: 1918: 1908: 1884: 1875: 1868: 1862: 1861: 1838:Pigment Cell Res 1833: 1827: 1826: 1806: 1797: 1794: 1788: 1787: 1759: 1739: 1728: 1727: 1707: 1696: 1695: 1685: 1661: 1655: 1654: 1623:Pigment Cell Res 1618: 1592:Human skin color 1544:structural color 1493:Impact on vision 1348: 1341: 1337: 1334: 1328: 1305: 1297: 1283:in photographs. 1277:pupillary reflex 1194:Wilson's disease 1188:Wilson's disease 1174:Wilson's disease 1139:Elizabeth Taylor 1005:white population 992: 976: 962: 942: 808:structural color 664: 661: 655: 639: 638: 631: 534:great horned owl 360:A 1997 study of 312:Associated with 296: 295: 246:common in human 244:hypopigmentation 110:structural color 73:of light by the 55:phenotypic trait 21: 5264: 5263: 5259: 5258: 5257: 5255: 5254: 5253: 5244:Facial features 5224: 5223: 5206: 5201: 5200: 5194:Wayback Machine 5185: 5181: 5158: 5154: 5144: 5142: 5118: 5114: 5087: 5083: 5078: 5074: 5064: 5062: 5053: 5052: 5048: 5039: 5035: 5004: 4997: 4950: 4946: 4899: 4895: 4848: 4841: 4831: 4829: 4811: 4804: 4767: 4763: 4719: 4713: 4709: 4678: 4674: 4663: 4659: 4632:Surv Ophthalmol 4628: 4624: 4611: 4607: 4594: 4590: 4583: 4561: 4557: 4550: 4528: 4521: 4476: 4472: 4441: 4437: 4398: 4394: 4363: 4359: 4320:Guymer, Robyn H 4316: 4312: 4282: 4278: 4233: 4229: 4219: 4217: 4215: 4185: 4181: 4171: 4169: 4154: 4150: 4140: 4138: 4121: 4117: 4111:Wayback Machine 4102: 4098: 4088: 4086: 4077: 4076: 4069: 4059: 4057: 4049: 4048: 4044: 4031: 4027: 4014: 4013: 4009: 3999: 3997: 3982: 3975: 3930: 3926: 3907: 3903: 3893: 3891: 3873: 3869: 3863:Blue eyed Koala 3861: 3857: 3844: 3843: 3839: 3832: 3814: 3810: 3753: 3749: 3739: 3737: 3722: 3718: 3678: 3674: 3663: 3661: 3658:The Independent 3652: 3651: 3647: 3637: 3635: 3620: 3613: 3606: 3602: 3565:Br J Ophthalmol 3557: 3553: 3547:Wayback Machine 3538: 3534: 3524: 3522: 3492:Nature Genetics 3484: 3477: 3469: 3465: 3459:Wayback Machine 3450: 3446: 3402: 3398: 3388: 3372: 3368: 3359: 3358: 3354: 3303: 3296: 3265: 3261: 3222: 3218: 3181: 3174: 3137: 3130: 3093: 3089: 3054: 3047: 3010: 3006: 2965: 2958: 2917:(9): e1004572. 2902: 2898: 2888: 2886: 2878: 2877: 2873: 2838: 2831: 2796:Cell Tissue Res 2792: 2788: 2757: 2753: 2747:Wayback Machine 2738:Larry Bickford 2737: 2733: 2727:Wayback Machine 2718: 2714: 2703: 2699: 2692: 2676: 2669: 2659: 2657: 2647: 2638: 2629: 2625: 2617: 2613: 2605: 2596: 2565: 2561: 2528:(5): e1000934. 2513: 2509: 2480:(12): 1443–52. 2469: 2456: 2411: 2407: 2353: 2346: 2276: 2272: 2261: 2259: 2257: 2241: 2237: 2186: 2175: 2145:Current Biology 2132: 2128: 2114: 2110: 2071: 2064: 2056: 2025: 2019: 2012: 1967: 1960: 1950: 1948: 1939: 1938: 1934: 1926: 1922: 1885: 1878: 1869: 1865: 1834: 1830: 1807: 1800: 1795: 1791: 1776: 1740: 1731: 1724: 1708: 1699: 1662: 1658: 1619: 1615: 1610: 1583: 1563:Corucia zebrata 1536:Bruno K Schultz 1508: 1495: 1470: 1464: 1389:or acquired by 1371:Heterochromia ( 1349: 1338: 1332: 1329: 1318: 1306: 1295: 1289: 1272:eye examination 1259: 1248: 1242: 1218: 1206:Golgi apparatus 1176: 1148: 1108: 1098:As a result of 1085: 1080: 1049:Aurès Mountains 1018: 999: 996:Sclater's lemur 993: 984: 977: 968: 963: 954: 943: 915:southern Europe 830:polygenic trait 761: 754: 665: 659: 656: 649: 640: 636: 629: 570: 540:within certain 511: 461:, and parts of 395: 388: 383: 381:Eye color range 362:White Americans 358: 167: 148:recessive trait 106:vitreous humour 35: 28: 23: 22: 15: 12: 11: 5: 5262: 5252: 5251: 5246: 5241: 5236: 5222: 5221: 5216: 5205: 5204:External links 5202: 5199: 5198: 5179: 5152: 5112: 5101:(3): 183–218. 5081: 5072: 5046: 5033: 4995: 4960:(2): 455–460. 4944: 4893: 4839: 4802: 4761: 4730:(2): 137–146. 4707: 4672: 4657: 4622: 4605: 4588: 4581: 4555: 4548: 4519: 4490:(884): 375–8. 4484:Postgrad Med J 4470: 4451:(2): 237–238. 4435: 4408:(5): 293–302. 4392: 4357: 4310: 4276: 4227: 4213: 4179: 4148: 4115: 4096: 4067: 4042: 4025: 4007: 3973: 3924: 3901: 3867: 3855: 3837: 3830: 3808: 3747: 3716: 3672: 3645: 3634:on 1 July 2013 3611: 3600: 3551: 3532: 3475: 3463: 3444: 3413:(3): 395–410. 3407:Human Genetics 3396: 3386: 3366: 3352: 3317:(4): 686–689. 3294: 3259: 3226:Doc Ophthalmol 3216: 3172: 3128: 3087: 3045: 3004: 2983:(2): 197–210. 2956: 2896: 2871: 2829: 2786: 2751: 2731: 2712: 2697: 2690: 2667: 2636: 2623: 2611: 2609:, p. 660. 2594: 2559: 2507: 2454: 2405: 2367:(9): 864–875. 2344: 2270: 2255: 2235: 2173: 2126: 2124:14 March 2009. 2108: 2075:Ann. Hum. Biol 2062: 2010: 1987:10.1086/510885 1958: 1947:on 5 June 2011 1932: 1920: 1876: 1863: 1828: 1817:(5): 498–501. 1798: 1789: 1774: 1744:Sing Bing Kang 1729: 1722: 1697: 1656: 1612: 1611: 1609: 1606: 1605: 1604: 1599: 1594: 1589: 1582: 1579: 1507: 1504: 1494: 1491: 1483:corneal limbus 1466:Main article: 1463: 1460: 1448:X-inactivation 1351: 1350: 1309: 1307: 1300: 1291:Main article: 1288: 1285: 1281:red eye effect 1258: 1255: 1244:Main article: 1241: 1238: 1217: 1214: 1175: 1172: 1164:uveal melanoma 1147: 1144: 1132:red-eye effect 1107: 1106:Red and violet 1104: 1084: 1081: 1079: 1076: 1035:Mie scattering 1017: 1014: 1001: 1000: 994: 987: 985: 978: 971: 969: 964: 957: 955: 944: 937: 871:Human Genetics 753: 750: 698:Central Europe 667: 666: 643: 641: 634: 628: 625: 617:United Kingdom 569: 566: 542:chromatophores 510: 507: 487:Southeast Asia 427:Southeast Asia 387: 384: 382: 379: 357: 354: 348: 347: 344: 338: 337: 334: 328: 327: 324: 318: 317: 310: 304: 303: 300: 166: 163: 77:medium in the 26: 9: 6: 4: 3: 2: 5261: 5250: 5247: 5245: 5242: 5240: 5237: 5235: 5232: 5231: 5229: 5220: 5217: 5215: 5211: 5208: 5207: 5195: 5191: 5188: 5183: 5175: 5171: 5167: 5163: 5156: 5141: 5137: 5134:(8): 1592–8. 5133: 5129: 5128: 5123: 5116: 5108: 5104: 5100: 5096: 5092: 5085: 5076: 5060: 5056: 5050: 5043: 5037: 5029: 5025: 5021: 5017: 5014:(2): 103–10. 5013: 5009: 5002: 5000: 4991: 4987: 4983: 4979: 4975: 4971: 4967: 4963: 4959: 4955: 4948: 4940: 4936: 4932: 4928: 4924: 4920: 4916: 4912: 4908: 4904: 4897: 4889: 4885: 4881: 4877: 4873: 4869: 4865: 4861: 4857: 4853: 4846: 4844: 4828: 4824: 4820: 4816: 4809: 4807: 4798: 4794: 4789: 4784: 4780: 4776: 4772: 4765: 4757: 4753: 4748: 4743: 4738: 4733: 4729: 4725: 4718: 4711: 4703: 4699: 4695: 4691: 4688:(3): 114–23. 4687: 4683: 4676: 4670: 4666: 4661: 4653: 4649: 4645: 4641: 4637: 4633: 4626: 4620: 4616: 4615: 4609: 4603: 4599: 4598: 4592: 4584: 4578: 4574: 4570: 4566: 4559: 4551: 4545: 4541: 4537: 4533: 4526: 4524: 4515: 4511: 4506: 4501: 4497: 4493: 4489: 4485: 4481: 4474: 4466: 4462: 4458: 4454: 4450: 4446: 4439: 4431: 4427: 4423: 4419: 4415: 4411: 4407: 4403: 4396: 4388: 4384: 4380: 4376: 4373:(2): 247–52. 4372: 4368: 4361: 4353: 4349: 4345: 4341: 4337: 4333: 4329: 4325: 4321: 4314: 4304: 4299: 4295: 4291: 4287: 4280: 4272: 4268: 4264: 4260: 4255: 4250: 4246: 4242: 4238: 4231: 4216: 4210: 4206: 4202: 4198: 4194: 4190: 4183: 4167: 4163: 4159: 4152: 4136: 4132: 4131: 4126: 4119: 4112: 4108: 4105: 4100: 4084: 4080: 4074: 4072: 4056: 4052: 4046: 4040: 4036: 4035: 4029: 4023: 4019: 4011: 3995: 3991: 3987: 3980: 3978: 3969: 3965: 3960: 3955: 3951: 3947: 3943: 3939: 3938:Croat. Med. J 3935: 3928: 3920: 3916: 3912: 3905: 3890: 3886: 3882: 3878: 3871: 3864: 3859: 3851: 3847: 3841: 3833: 3827: 3822: 3821: 3812: 3805: 3800: 3796: 3791: 3786: 3782: 3778: 3774: 3770: 3766: 3762: 3758: 3751: 3735: 3731: 3727: 3720: 3712: 3708: 3704: 3700: 3696: 3692: 3689:(2): 177–87. 3688: 3684: 3676: 3659: 3655: 3649: 3633: 3629: 3625: 3618: 3616: 3609: 3604: 3596: 3592: 3587: 3582: 3578: 3574: 3571:(7): 549–52. 3570: 3566: 3562: 3555: 3548: 3544: 3541: 3536: 3521: 3517: 3513: 3509: 3505: 3501: 3497: 3493: 3489: 3482: 3480: 3472: 3467: 3460: 3456: 3453: 3448: 3441: 3436: 3432: 3428: 3424: 3420: 3416: 3412: 3408: 3400: 3393: 3389: 3387:9780091013707 3383: 3379: 3378: 3370: 3362: 3356: 3348: 3344: 3340: 3336: 3332: 3328: 3324: 3320: 3316: 3312: 3308: 3301: 3299: 3290: 3286: 3282: 3278: 3274: 3270: 3263: 3255: 3251: 3247: 3243: 3239: 3235: 3232:(4): 367–76. 3231: 3227: 3220: 3212: 3208: 3203: 3198: 3194: 3190: 3186: 3179: 3177: 3168: 3164: 3159: 3154: 3150: 3146: 3142: 3135: 3133: 3124: 3120: 3115: 3110: 3107:(6): 853–60. 3106: 3102: 3098: 3091: 3083: 3079: 3075: 3071: 3067: 3063: 3059: 3052: 3050: 3041: 3037: 3032: 3027: 3023: 3019: 3015: 3008: 3000: 2996: 2991: 2986: 2982: 2978: 2974: 2970: 2963: 2961: 2952: 2948: 2943: 2938: 2934: 2930: 2925: 2920: 2916: 2912: 2911:PLOS Genetics 2908: 2900: 2885: 2881: 2875: 2867: 2863: 2859: 2855: 2851: 2847: 2843: 2836: 2834: 2825: 2821: 2817: 2813: 2809: 2805: 2802:(2): 387–95. 2801: 2797: 2790: 2782: 2778: 2774: 2770: 2766: 2762: 2755: 2748: 2744: 2741: 2735: 2728: 2724: 2721: 2716: 2710: 2706: 2701: 2693: 2687: 2683: 2682: 2674: 2672: 2656: 2652: 2645: 2643: 2641: 2633: 2627: 2620: 2615: 2608: 2603: 2601: 2599: 2590: 2586: 2582: 2578: 2575:(5): 659–63. 2574: 2570: 2563: 2555: 2551: 2546: 2541: 2536: 2531: 2527: 2523: 2522:PLOS Genetics 2519: 2511: 2503: 2499: 2495: 2491: 2487: 2483: 2479: 2475: 2467: 2465: 2463: 2461: 2459: 2450: 2446: 2441: 2436: 2432: 2428: 2425:(2): 411–23. 2424: 2420: 2416: 2409: 2400: 2396: 2392: 2388: 2384: 2380: 2375: 2370: 2366: 2362: 2358: 2351: 2349: 2341: 2336: 2332: 2327: 2322: 2318: 2314: 2309: 2304: 2299: 2294: 2290: 2286: 2282: 2274: 2258: 2252: 2248: 2247: 2239: 2230: 2226: 2222: 2218: 2213: 2208: 2204: 2200: 2196: 2192: 2184: 2182: 2180: 2178: 2169: 2165: 2160: 2155: 2151: 2147: 2146: 2141: 2137: 2130: 2123: 2122: 2121:New Scientist 2117: 2112: 2104: 2100: 2096: 2092: 2088: 2084: 2081:(6): 657–66. 2080: 2076: 2069: 2067: 2055: 2051: 2047: 2043: 2039: 2036:(8): 327–32. 2035: 2031: 2024: 2017: 2015: 2006: 2002: 1997: 1992: 1988: 1984: 1981:(2): 241–52. 1980: 1976: 1972: 1965: 1963: 1946: 1942: 1936: 1929: 1924: 1916: 1912: 1907: 1902: 1898: 1894: 1890: 1883: 1881: 1873: 1867: 1859: 1855: 1851: 1847: 1844:(2): 129–31. 1843: 1839: 1832: 1824: 1820: 1816: 1812: 1805: 1803: 1793: 1785: 1781: 1777: 1771: 1767: 1763: 1758: 1757:10.1.1.87.418 1753: 1749: 1745: 1738: 1736: 1734: 1725: 1719: 1715: 1714: 1706: 1704: 1702: 1693: 1689: 1684: 1679: 1675: 1671: 1667: 1660: 1652: 1648: 1644: 1640: 1636: 1632: 1629:(6): 454–64. 1628: 1624: 1617: 1613: 1603: 1600: 1598: 1595: 1593: 1590: 1588: 1585: 1584: 1578: 1576: 1571: 1569: 1565: 1564: 1559: 1556: 1551: 1549: 1545: 1539: 1537: 1533: 1532:Rudolf Martin 1529: 1525: 1521: 1517: 1512: 1503: 1501: 1500:visual acuity 1490: 1488: 1484: 1474: 1469: 1459: 1457: 1453: 1449: 1444: 1442: 1437: 1435: 1430: 1429:Lee Van Cleef 1425: 1423: 1419: 1414: 1412: 1408: 1404: 1400: 1396: 1392: 1388: 1384: 1380: 1376: 1375: 1365: 1357: 1347: 1344: 1336: 1333:November 2012 1326: 1322: 1316: 1315: 1310:This section 1308: 1304: 1299: 1298: 1294: 1287:Heterochromia 1284: 1282: 1278: 1273: 1269: 1265: 1254: 1252: 1247: 1237: 1235: 1231: 1227: 1223: 1213: 1211: 1207: 1203: 1199: 1195: 1189: 1185: 1180: 1171: 1169: 1165: 1161: 1157: 1152: 1143: 1140: 1135: 1133: 1129: 1125: 1121: 1112: 1103: 1101: 1094: 1089: 1078:Special cases 1075: 1072: 1070: 1066: 1062: 1058: 1054: 1050: 1046: 1045:Shawia people 1043: 1038: 1036: 1032: 1022: 1013: 1011: 1006: 997: 991: 986: 982: 975: 970: 967: 961: 956: 952: 948: 947:Birman kitten 941: 936: 935: 934: 932: 928: 924: 920: 916: 912: 907: 905: 901: 896: 894: 890: 886: 882: 881: 876: 872: 867: 865: 864:out of Africa 860: 856: 852: 848: 844: 843: 838: 833: 831: 827: 822: 820: 816: 811: 809: 805: 801: 797: 793: 789: 785: 784:vitreous body 776: 770: 765: 759: 746: 741: 733: 729: 727: 721: 719: 715: 711: 707: 703: 699: 695: 691: 686: 684: 680: 676: 675: 663: 653: 647: 644:This section 642: 633: 632: 624: 622: 618: 614: 610: 605: 603: 599: 594: 591: 582: 574: 565: 563: 562:South America 559: 555: 551: 547: 543: 539: 535: 531: 526: 524: 515: 502: 498: 496: 492: 488: 484: 480: 476: 472: 468: 464: 460: 456: 452: 448: 444: 440: 436: 432: 428: 424: 420: 411: 407: 399: 393: 378: 376: 371: 368: 365: 363: 353: 345: 343: 340: 339: 335: 333: 330: 329: 325: 323: 320: 319: 315: 311: 309: 306: 305: 301: 298: 297: 294: 292: 288: 284: 280: 276: 273: 269: 265: 261: 257: 253: 249: 245: 241: 237: 232: 230: 229:chromosome 15 226: 225: 220: 219: 213: 211: 210:out of Africa 207: 203: 199: 194: 192: 188: 182: 180: 176: 172: 162: 160: 157: 153: 149: 145: 144: 139: 138: 132: 130: 126: 122: 118: 113: 111: 107: 103: 99: 95: 91: 87: 82: 80: 76: 72: 68: 64: 60: 56: 53: 49: 41: 37: 33: 19: 5182: 5174:the original 5169: 5165: 5155: 5143:. Retrieved 5131: 5125: 5115: 5098: 5094: 5084: 5075: 5063:. Retrieved 5059:the original 5049: 5036: 5011: 5007: 4957: 4953: 4947: 4906: 4902: 4896: 4858:(1): 91–95. 4855: 4851: 4830:. Retrieved 4818: 4781:(1): 80–91. 4778: 4774: 4764: 4727: 4723: 4710: 4685: 4681: 4675: 4660: 4635: 4631: 4625: 4612: 4608: 4595: 4591: 4564: 4558: 4531: 4487: 4483: 4473: 4448: 4444: 4438: 4405: 4401: 4395: 4370: 4366: 4360: 4330:(6): 465–9. 4327: 4323: 4313: 4293: 4289: 4279: 4244: 4240: 4230: 4218:. Retrieved 4196: 4192: 4182: 4170:. Retrieved 4166:the original 4161: 4151: 4139:. Retrieved 4135:the original 4128: 4118: 4099: 4087:. Retrieved 4082: 4058:. Retrieved 4054: 4045: 4032: 4028: 4021: 4010: 3998:. Retrieved 3994:the original 3989: 3944:(4): 381–6. 3941: 3937: 3927: 3919:the original 3914: 3904: 3892:. Retrieved 3880: 3870: 3858: 3850:the original 3840: 3819: 3811: 3802: 3764: 3760: 3750: 3738:. Retrieved 3734:the original 3729: 3719: 3686: 3682: 3675: 3662:. Retrieved 3657: 3648: 3636:. Retrieved 3632:the original 3603: 3568: 3564: 3554: 3535: 3523:. Retrieved 3495: 3491: 3466: 3447: 3438: 3410: 3406: 3399: 3391: 3376: 3369: 3355: 3314: 3310: 3275:(3): 293–7. 3272: 3269:Exp. Eye Res 3268: 3262: 3229: 3225: 3219: 3195:(6): 811–4. 3192: 3188: 3148: 3144: 3104: 3100: 3090: 3068:(3): 384–6. 3065: 3061: 3021: 3017: 3007: 2980: 2976: 2914: 2910: 2899: 2887:. Retrieved 2883: 2880:"Amber eyes" 2874: 2849: 2845: 2799: 2795: 2789: 2767:(3): 202–8. 2764: 2760: 2754: 2734: 2715: 2700: 2680: 2658:. Retrieved 2654: 2626: 2614: 2572: 2568: 2562: 2525: 2521: 2510: 2477: 2473: 2422: 2418: 2408: 2364: 2360: 2338: 2288: 2284: 2273: 2260:. Retrieved 2245: 2238: 2194: 2190: 2149: 2143: 2129: 2119: 2111: 2078: 2074: 2054:the original 2033: 2030:Trends Genet 2029: 1978: 1974: 1949:. Retrieved 1945:the original 1935: 1923: 1896: 1892: 1870:Morris, PJ. 1866: 1841: 1837: 1831: 1814: 1810: 1792: 1747: 1712: 1676:(3): 293–9. 1673: 1670:Exp. 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