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Mackey 2011

This document reviews the classification of iris color. It discusses factors that influence iris color like genetics, environment, and skin and hair pigmentation. It presents a nine category classification system for iris color ranging from light to dark blue, green, brown, and mixtures of colors. The classification aims to standardize descriptions of iris color which may provide insights into eye disease and conditions that affect color.

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0% found this document useful (0 votes)
40 views10 pages

Mackey 2011

This document reviews the classification of iris color. It discusses factors that influence iris color like genetics, environment, and skin and hair pigmentation. It presents a nine category classification system for iris color ranging from light to dark blue, green, brown, and mixtures of colors. The classification aims to standardize descriptions of iris color which may provide insights into eye disease and conditions that affect color.

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catherin
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We take content rights seriously. If you suspect this is your content, claim it here.
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Clinical and Experimental Ophthalmology 2011; 39: 462–471 doi: 10.1111/j.1442-9071.2010.02487.

Review

Classification of iris colour: review and refinement


of a classification schema
David A Mackey MD FRANZCO,1,2,3 Colleen H Wilkinson BOrth(Hons),2 Lisa S Kearns BOrth&OphthalSci(Hons)2
and Alex W Hewitt MBBS PhD2
1
Lions Eye Institute, Centre for Ophthalmology and Visual Science, University of Western Australia, Perth, Western Australia, 2Centre
for Eye Research Australia, University of Melbourne, Department of Ophthalmology, Royal Victorian Eye and Ear Hospital, Melbourne,
Victoria and 3Department of Ophthalmology, Royal Hobart Hospital, University of Tasmania, Hobart, Tasmania, Australia

ABSTRACT colour will result in more standardized classification of


iris colour and investigation of its role in eye disease.
Eye colour or, more accurately, iris colour is one of the
most obvious physical characteristics of a person. Key words: albinism, eye colour classification, hetero-
European parents frequently ask the colour of their chromia, iris colour, pigmentation.
newborn’s eyes, only to see the iris change dramati-
cally during their child’s first year of life. Genetic and
epidemiological findings have uncovered further VARIATION IN EYE COLOUR
details about the basis for iris colour, which may have Environmental factors, particularly sunlight, have
important implications for further research and treat- driven biological variation between populations
ment of some eye diseases and ocular characteristics. living at different latitudes. People living at lower
Surprisingly there is no widely recognized classifica- latitudes where the ultra violet radiation (UVR)
tion system for eye colour. An added difficulty when intensity is high generally have darker skin, hair and
trying to devise an international system is that subtle eye pigmentation that act as a protective barrier
against the harmful effects of UVR. In contrast,
differences in colour description exist between lan-
people living at higher latitudes tend to have fairer
guages (e.g. hazel vs. auburn). We reviewed the recent skin, hair and eyes to facilitate vitamin D production
and very early literature pertaining to eye colour in lower levels of UVR.1 Modern migration of Euro-
classification. Recent genetic investigations of eye peans to new world countries and, to a lesser extent,
colour have tended to either use simple (three- Africans and Asians to European countries has
category grading systems) or more complex digital resulted in people living in environments that differ
colour grading. We present a nine-category grading from those in which they evolved over the last
system. Categories in this novel schema include: (i) 100 000 years.
light blue; (ii) darker blue; (iii) blue with brown
peripupillary ring; (iv) green; (v) green with brown iris FACTORS INFLUENCING EYE COLOUR
ring; (vi) peripheral green central brown; (vii) brown
with some peripheral green; (viii) brown; and (ix) dark Colour is a combination of hue, saturation and
brightness. It is influenced by the centre surround
brown. Although different observers may categorize a
phenomenon and, in the case of eye colour, the per-
person’s eye colour differently, it is generally only by
son’s skin and hair colour (eyelashes and eyebrows),
an adjacent category. We also describe a continuum of their conjunctival colour (white or inflamed) and a
iris pigmentation from a small ring of brown around delineated limbus (as in the same phenomenon as
the pupil to almost complete brown with small periph- dark outlines around colours on the television
eral flecks. Digital publishing and assessment of iris screen). The cornea can influence iris colour

䊏 Correspondence: Professor David A Mackey, Lions Eye Institute, Centre for Ophthalmology and Visual Science, University of Western Australia, 2
Verdun St, Nedlands, WA 6009, Australia. Email: d.mackey@utas.edu.au
Received 24 August 2010; accepted 25 October 2010.

© 2011 The Authors


Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
Eye colour classification 463

Table 1. Conditions that affect iris colour8 Table 2. Genes associated with eye colour7,10

Congenital Acquired Gene Gene map locus OMIM number


Waardenburg’s Fuchs’ heterochromic iridocyclitis HERC2/OCA2 15q11.2-q12 611409
syndrome SLC2A4 17p13 138190
Horner’s syndrome Treatment with latanoprost (Xalatan), TYR 11q14-q21 606933
bimatoprost (Lumigan), travaprost TYRP1 9p23 115501
(Travatan) for glaucoma SLC45A2 5p13.3 606202
Aniridia Trauma (including sideroris)/surgery IRF4 6p25-p23 601900
Coloboma Iris melanoma DSCR9 21q22.13 Recent publication, not
Sturge–Weber on OMIM7
syndrome LYST 1q42.3 Recent publication, not
on OMIM7
NPLOC4 17q25.3 Recent publication, not
appearance. For example, corneal opacities, corneal on OMIM7
thickness, corneal curvature and anterior chamber KITLG 12q21.33 184745
depth all affect the way the light in the eye is MC1R 16q24.3 155555
reflected. The external lighting and direction of illu-
mination will influence this; notably commercial
photographers will highlight the colour of a model’s eyes to have increased pigment around the pupil
eyes by using side lighting. The crypts, furrows and affect colour. Iris transillumination (providing a
collarettes of the iris will also influence eye colour.2 lighter colour) is frequently observed in albinism,
In some cases the black posterior iris is visible at the and occasionally in eyes with blue irides, following
pupil margin. The dilation of the pupil itself will trauma, pigment dispersion syndrome or herpetic
influence the colour of the iris, particularly where infections.
there is more peripupillary pigmentation. Post-
mortem studies have shown that brown irises have
40% more melanin than other colours.3 The number GENES FOR EYE COLOUR
of melanocytes, the proportion of melanocytes and Eye colour has played an important role in under-
iris stromal cellularity are not major contributors to standing genetics, although a dominant–recessive
iris colour,4 although the average melanosomal area model for iris colour is overly simplistic for this
per perinuclear cytoplasmic area (AMAC) and polygenic trait. The concept of inheritance is first
average number of melanosomes per perinuclear understood by many people through comments such
area (AMNC) within superficial melanocytes differ as ‘he has eyes like his father’. The simple model that
significantly across iris colour groups.5 Interestingly brown eyes are dominant and blue eyes are recessive
eye colour has the potential to change over time. is demonstrated in many but not all families and the
Many Caucasian children are born with blue eyes uncommon and potentially embarrassing instance of
and their eye pigment often increases, predomi- brown-eyed children of blue-eyed parents needs
nantly during their first year of life although this careful explanation of heterogeneity. Indeed this
process may continue for up to 6 years or beyond. occurrence inspired the career of one of the greatest
There are surprisingly few data on long-term geneticists, Victor McKussick, who like his identical
changes in eye colour through adolescence and adult twin was the brown-eyed son of blue-eyed parents.9
life.6 A large cross-sectional study revealed that age Hair, skin and eye colour are quite evidently linked
and gender seem to have a significant effect on eye in albinism; however, genome-wide association
colour.7 scans to date do not support the premise that the
same genes are involved in the subtle variability in
eye and skin colour in Europeans. Indeed recent
EYE DISEASES RELATED TO EYE COLOUR
genome-wide association scans have identified a
Heterochromia iridis is a difference in pigmentation series of genes involved in eye colour variation in
of the iris between a person’s eyes and may be inher- people of Northern European ancestry (Table 2).7,10 A
ited or acquired by disease or injury (common causes single base change (rs12913832; T to C) within
are listed in Table 1).8 Specific disease-associated intron 86 of the upstream HERC2 locus explains
features such as Brushfield spots (white tufts seen almost all of the association with blue-brown eye
more often in individuals with Down syndrome) can colour.11–13 It is thought that this single nucleotide
cause the eye to have a lighter colour. Conversely, the polymorphism acts as part of a highly evolutionary
distribution of naevi, other pigmented lesions such conserved regulatory element required for OCA2
as Lisch nodules (as seen in Neurofibromatosis Type gene activation through chromatin remodelling.11
1) or melanosis and the tendency of hazel-coloured Although there is one major gene for eye colour, the
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
464 Mackey et al.

remaining genetic influences are due to a large col- colours using a mixture of pigments, such as dry
lection of genes, each with much less effect. With ‘fresco’ pigments: cadium red, titanium white, terre
the identification of the single nucleotide polymor- de Sienne ocre, burned bones black, cobalt blue,
phisms associated with different eye colours it is yellow (P Loyer, pers. comm., 2010). Swiss Anthro-
becoming possible for forensic scientists to infer the pologist Rudolf Martin, who was Professor of Physi-
eye colour of suspects based on DNA evidence.7 In cal Anthropology at the University of Zurich, made
addition historic DNA samples can be used to iden- the first attempt at using a standard set of artificial
tify eye colour of famous people long dead, such as eyes to classify eye colour. In 1903 he developed the
the astronomer Copernicus.14 ‘Augenfarbentafel’ or eye colour chart, which con-
sisted of 16 glass eyes progressively arranged in
shades from dark brown (number 1) to light blue
STANDARDIZATION OF EYE COLOUR
(number 16) (Fig. 1).23 When observers differed in
There have been various attempts to develop a uni- their classifications, it was usually only by adjacent
versal classification system for eye colour. The earli- numbers.
est study on eye colour was by Pétrequin in 1843,15 Problems with the consistency of results from
who used categories of grey, blue, auburn, brown, classification with the Martin’s eyes led to the devel-
black. Wilde (Oscar’s father)16 described grey, blue, opment of new classification systems.21 In 1942,
hazel and brown. Cornaz17 described blue (with grey, Riddell developed a descriptive classification with
blue and green tints) and brown (with yellow, hazel, 1 = blue, 2 = grey, 3 = green, 4 = yellow, 5 = tan and
auburn, light brown, dark brown and black tints). 6 = chocolate.24–26 An eye’s colour is classified as a
Galton, the famous geneticist, classified eye colour three-digit number, with the first digit describing the
under eight headings: 1, light blue; 2, blue, dark general colour of the iris, the second digit the diffuse
blue; 3, grey, blue-green; 4, dark grey, hazel; 5, light pigmentation (which is 0 if none) and the third digit
brown; 6, brown; 7, dark brown; 8, black, for his the discrete spots of pigment, which may also be 0.
research into the ‘Family likeness in eye-colour’ but In 1946, Grieve and Morant21 developed a 14-
simplified this to three headings: light, hazel (=inter- category classification: no brown (A = light blue,
mediate tints) and dark.18 Tocher19 delineated blue, B = light grey, C = dark blue, D = dark grey); more
grey, heterochromic mixed and brown. Brownlee20 blue or grey than brown (E = trace of brown,
described four groups: ‘(1) pure blue eyes in which F = more brown, G = still more brown, H = most
there is no pigment visible on the iris, such grey as brown); more brown than blue or grey (I = least
appears being due to strands of connective tissue; (2) brown, J = more brown, K = most brown); ‘pure’
grey or pale yellow, in which there is always visible brown (L = light, M = medium, N = dark).
pigment present in little masses quite distinct from The use of colour photography to classify eye
definite strands of connective tissue; (3) the deep colour did not progress because of the variability in
yellow eye, more or less rare form, not much exceed- developing and printing of images and the lack of
ing 1% of the adult population as seen in Glasgow; stability over time with prints and slides fading or
(4) the dark brown or chocolate eye, of which the yellowing. Thus even until the 1990s researchers
shades vary, but in all of which the iris is sensibly were using comparisons with standardized glass
the same colour from the inner margin to the outer’. eyes.6 The latest methods involve scanned digital
All four of these can be mixed with each other giving images that are then read against standardized
10 groups. computer colours for hue and saturation.7 With the
The lack of standardization of eye colour with no advent of large population-based eye studies in the
widely accepted classifications has been due to the 1990s, the Iris Colour Classification System was
fact that colour print illustrations could fade with developed (Fig. 2).27 This system has avoided inher-
time. The earliest published standard was based on ent problems associated with heterogeneity of
painted artificial eyes.21 Although artificial eyes were definitions. Four standard photos were used for
made to decorate statues for millennia, the first pros- various large epidemiological studies. Grade 1: blue
theses were made of glass in Venice. It is Shakes- or gray iris with brown or yellow specks equal to (in
peare in 1606 who first mentions a glass eye, in Act approximate percentage of total iris area) or less than
IV, Scene 6 of King Lear:22 ‘Get thee glass eyes/And, in standard A. Grade 2: blue, gray or green iris with
like a scurvy politician seem/To see the things thou brown or yellow specks equal to or less than in
dost not’ (King Lear to the Earl of Gloucester). Glass standard B but greater than in standard A. Grade 3:
eyes changed over time with the development of green or light brown iris with brown or yellow
new materials. Ocularists would have a collection specks equal to or less than in standard C but greater
of different coloured eyes to match the remaining eye than in standard B. Grade 4: brown iris with colour
of a person who had lost an eye, usually from injury. equal to or less than in standard D but greater than
Modern ocularists can reproduce most human eye in standard C. Grade 5: brown iris, darker than
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
Eye colour classification 465

Figure 1. Set of 16 Martin’s eyes, graded from the darkest brown (number 1) to a light blue (number 16) and the eyes in this photograph
are not in the correct order. (Reproduced with permission: Collection, Powerhouse Museum, Sydney. Photo: Sotha Bourn.)

Figure 2. Iris Colour Classifica-


tion System: A, B, C, D. (Repro-
duced with permission: Invest
Ophthalmol Vis Sci and the
authors.27)

Standard A Standard C

Standard B Standard D

standard D. Although the grading system has five change in iris colour.28,29 Frudakis et al. attempted
categories, most subsequent studies have generally to objectify the quantification of iris colour using
restricted to four categories. digital spectroscopy to accurately predict iris
Subsequent recognition that the new prostaglan- melanin content.30 An iris colour score (C) was deter-
din analogues cause increased pigmentation of some mined using a function incorporating the basic ele-
eyes (usually darkening partly brown eyes) led to ments of the C.I.E. colour wheel (luminosity, red
some other digital and computer-based eye colour reflectance, blue reflectance, green reflectance). Iris
classifications of red/green/blue that could detect colour scores ranged from 2.9 to 0.9 (average = 2.07).
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
466 Mackey et al.

Figure 3. Blue, hazel/green or


brown.

The distribution of iris colour scores was continuous METHOD OF IRIS COLOUR GRADING
but bimodal with one peak around 1.85 (‘brown’
iris) and another around 2.65 (‘blue’ iris). They A grader sorted digital images of the face photos
noted that blue irides often have peripupillary rings containing both eyes into colour bin folders and then
or sectors of brown, red or yellow and many irides re-sorted each bin moving up or down a colour
that appear to be of a light colour from a distance are category. This was sorted by a first grader (DAM),
revealed upon closer inspection as a mix of light and second grader (CHW) and again by the first grader
dark regions. Wilkerson et al. in 19964 and Imesch (DAM). Many photographs could easily be moved
et al. in 19965 emphasized the presence of a brown back and forth between adjacent categories but
peripupillary ring, as reviewed by Sturm et al.11 To rarely beyond the adjacent category. Final colour was
their three main colours: uniform blue, uniform then coded with the main twin database.
hazel or uniform brown were added those with a In cases where photographs of the iris with
darker peripupillary zone: blue ring, hazel ring or dilated and undilated pupils were available, the
brown ring. A 2008 paper proposed a 24-scale clas- undilated photograph was used. To observe the
sification from 1 (least pigmentation) to 24 (most effect of dilation of the pupil on eye colour, we
pigmentation), which is very similar to the earlier selected 15 randomly chosen individuals who had
Martin’s eyes.31 Despite all the subclassification, photos before and after dilation. In the recent study
much successful genetic research work has been some participants had been dilated although others
done by classifying eyes or photos of eyes simply as were not.7 Eleven individuals (73.3%) had no
blue, hazel/green or brown (Fig. 3).32,33 change in eye colour. Of the four that changed, two
changed from blue with a brown ring to green with
a brown ring, one changed from blue with a brown
CATEGORIZATION OF IRIS COLOUR FOR THE ring to green and one changed from mid hazel (half
brown) to slight peripheral green (Fig. 6). Thus
TWINS EYE STUDY IN TASMANIA dilation appears to make a blue + brown eye look
We wished to further investigate eye colour in the green and peripheral green is lost with dilation.
twins from the Twins Eye Study in Tasmania.34 We Although others have suggested the pupillary
refined the original three-category characterization of pigment ring as a separate entity4,5,29 we felt this
eye colour used in the Brisbane Adolescent Twins was more like a continuum from a narrow area of
Study33 to include intervening categories. As there is peripapillary pigment to almost completely brown
considerable subjective variation in classifying eye iris with some peripheral green flecks. This con-
colour, we aimed to ensure we had enough catego- tinuum is displayed across 12 eyes in Figure 7.
ries that subjects would not be moved more than two To establish the validity of the new classification
categories approximately 5% of the time. Blue/hazel- we used iris photographs obtained from an indepen-
green/brown became light, bright or uniform dent cohort as part of the Raine Eye Health Study.35
blue/darker or irregular blue/green with ring of Two orthoptists graded 360 colour iris photographs.
brown around pupil/green periphery and brown To reduce colour desaturation, each grader viewed a
central/mainly brown with some peripheral green/ white image between each colour iris classification.
total brown. Two other categories – uniform green Intra-grader reproducibility was assessed by the
and blue with ring of brown around iris – were also repeat grading of images after 1 week. Analyses
created (Fig. 4). The eight categories are shown in were performed using Stata/SE 10.0 (Stata Corp.,
Figure 5. A ninth category of dark brown (found in College Station, TX, USA). High intra- and inter-
Melanesian/East Asian and African eyes) is also grader reproducibility was found (k = 0.74 and 0.64,
possible but was not used in the Tasmanian respectively). Nine images (2.5%) were found to
cohort. have a colour classification disparity of three bins
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
Eye colour classification 467

Figure 4. Development of inter-


mediate categories from three
original groups.

between the two graders. In assessing intra-grader her makeup and hair. The historic problem of paint-
reproducibility, only two images (0.6%) were found ing and prints changing colour over time impeded a
to have a three-bin colour classification disparity. consistent reference source. Even now chemicals of
films or electronic camera sensors changing may
influence the collection of colour images and even
CONCLUSION though the digital classification systems seem robust,
the output of printing and monitors may change over
Although complex digital analysis of colour images
time.
is possible, a simple three-category classification was
highly successful in identifying the OCA2 gene as the
main gene for variation in eye colour in Europeans.
ACKNOWLEDGEMENTS
We have developed a broader classification of the
three categories, inserting intermediate categories, We would like to thank the participants in the Twins
noting that in hazel/green eyes there is a relative Eye Study in Tasmania and Raine Eye Health Study.
increase in pigment extending from the pupil margin We would like to thanks Sandra Oates and Seyhan
to the outer iris, with the peripheral iris being the Yazar (Lions Eye Institute, Perth, Western Australia,
last area to be pigmented. We did not observe people Australia) for allowing us to use their validation data
with a brown peripheral iris and a blue or green on checking eye colour. We thank the following
central iris. Iris naevi and Brushfield spots can com- organizations for their financial support: Clifford
plicate colour classification but dilated pupils seem Craig Medical Research Trust, Ophthalmic Research
to merge the iris colour and in our experience change Institute of Australia (ORIA), Peggy and Leslie
the classification. The features peripheral to the iris – Cranbourne Foundation, National Health and
limbus, sclera and vessels, lashes, lids margins and Medical Research Foundation Project Grant (2005–
skin colour – all influence the eye colour and are part 2007), Jack Brockhoff Foundation, Centre for Eye
of a billion dollar cosmetics industry. Colours such as Research Australia receives Operational Infrastruc-
grey and violet were not seen and in photographs ture Support from the Victorian Government. David
would appear to be blue. Elizabeth Taylor’s ‘violet’ Mackey is the recipient of the Pfizer Australia Senior
eyes are blue with surrounding black and red from Research Fellowship.
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
468

a b c d

e f g h

Figure 5. Randomly selected sample colours from Twins Eye Study in Tasmania: (a) light blue; (b) darker blue; (c) blue with brown pupil ring; (d) green; (e) green with brown iris
ring; (f) peripheral green central brown; (g) brown with some peripheral green; and (h) brown (dark brown not shown).

© 2011 The Authors


Mackey et al.

Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
Eye colour classification 469

Figure 6. Iris colour change after dilation. (a) Blue with brown ring to green with brown ring; (b) blue with brown ring to green; (c) mid
hazel to slight peripheral green; and (d) blue with brown ring to green with brown ring. Note the defocused images tend to give a more
uniform colour.
© 2011 The Authors
Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
470 Mackey et al.

REFERENCES
1. Jablonski NG. The evolution of human skin and skin
color [Review]. Annu Rev Anthropol 2004; 33: 585–623.
2. Larsson M, Pedersen NL. Genetic correlations among
texture characteristics in the human iris. Mol Vis 2004;
10: 821–31.
3. Wielgus AR, Sarna T. Melanin in human irides of
different color and age of donors. Pigment Cell Res 2005;
18: 454–64.
4. Wilkerson CL, Syed NA, Fisher MR, Robinson NL,
Wallow IH, Albert DM. Melanocytes and iris color.
Light microscopic findings. Arch Ophthalmol 1996; 114:
437–42.
5. Imesch PD, Bindley CD, Khademian Z et al. Melano-
cytes and iris color. Electron microscopic findings. Arch
Ophthalmol 1996; 114: 443–7.
Figure 7. Continuum of green with brown ranging from small pupil ring to almost complete brown with peripheral green flecks.
6. Bito LZ, Matheny A, Cruickshanks KJ, Nondahl DM,
Carino OB. Eye color changes past early childhood.
The Louisville Twin Study. Arch Ophthalmol 1997; 115:
659–63.
7. Liu F, Wollstein A, Hysi PG et al. Digital quantification
of human eye color highlights genetic association of
three new loci. PLoS Genet 2010; 6: e1000934.
8. Imesch PD, Wallow IH, Albert DM. The color of the
human eye: a review of morphologic correlates and of
some conditions that affect iridial pigmentation. Surv
Ophthalmol 1997; 41 (Suppl. 2): S117–23.
9. McKusick VA. Mendelian Inheritance in Man: A Catalog of
Human Genes and Genetic Disorders, 12th edn. Baltimore,
MD: Johns Hopkins University Press, 1998.
10. Sulem P, Gudbjartsson DF, Stacey SN et al. Genetic
determinants of hair, eye and skin pigmentation in
Europeans. Nat Genet 2007; 39: 1443–52.
11. Sturm RA, Duffy DL, Zhao ZZ et al. A single SNP in an
evolutionary conserved region within intron 86 of the
HERC2 gene determines human blue-brown eye color.
Am J Hum Genet 2008; 82: 424–31.
12. Eiberg H, Troelsen J, Nielsen M et al. 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 2008; 123: 177–87.
13. Sturm RA, Larsson M. Genetics of human iris colour
and patterns. Pigment Cell Melanoma Res 2009; 22: 544–
62.
14. Bogdanowicz W, Allen M, Branicki W, Lembring M,
Gajewska M, Kupiec T. Genetic identification of
putative remains of the famous astronomer Nico-
laus Copernicus. Proc Natl Acad Sci USA 2009; 106:
12279–82.
15. Pétrequin J. Sur les diverses couleurs de l’iris et leurs
proportions dans nos climats. Ann. d’oculistique 1843;
10: 120–5.
16. Wilde WR. An essay on the malformations and con-
genital diseases of the organs of sight. Dublin 1862.
17. Cornaz E. Recherches statistiques sur la fréquence
comparative des couleurs de l’iris. Ann. d’oculistique
1854; 31: 251–80.
18. Galton F. Family likeness in eye-colour. Proc R Soc Lond
1886; 40: 402–16.

© 2011 The Authors


Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists
Eye colour classification 471

19. Tocher JF. Pigmentation survey of school children in 29. Niggemann B, Weinbauer G, Vogel F, Korte R. A stan-
Scotland. Biometrika 1908; 6: 129–235. dardized approach for iris color determination. Int J
20. Brownlee J. On inheritance of hair and eye colour. Proc Toxicol 2003; 22: 49–51.
R Soc Edinb 1912; 32: 458–74. 30. Frudakis T, Terravainen T, Thomas M. Multilocus
21. Grieve J, Morant GM. Record of eye colours for British OCA2 genotypes specify human iris colors. Hum Genet
populations and a description of a new eye-colour 2007; 122: 311–26.
scale. Ann Eugen 1946; 13: 161–71. 31. Franssen LCJ, van den Berg TJ. Grading of Iris color
22. Shakespeare W. King Lear. 1606. Accessed July 2010. with an extended photographic reference set. J Optom
Available from: http://nfs.sparknotes.com/lear/page_ 2008; 1: 36–40.
244.html 32. Muinos Diaz Y, Saornil MA, Almaraz A, Munoz-
23. Oetteking B. Rudolf Martin. Am Anthropol 1926; 28: Moreno MF, Garcia C, Sanz R. Iris color: validation of
414–17. a new classification and distribution in a Spanish
24. Riddell WJB. Studies in the classification of eye colour population-based sample. Eur J Ophthalmol 2009; 19:
I. Ann Eugen 1942; 2: 245–59. 686–9.
25. Riddell WJB. Studies in the classification of eye colour 33. Zhu G, Evans DM, Duffy DL et al. A genome scan for
II. Ann Eugen 1943; 12: 24–30. eye color in 502 twin families: most variation is due to
26. Riddell WJB. Studies in the classification of eye colour a QTL on chromosome 15q. Twin Res 2004; 7: 197–210.
III. Ann Eugen 1945; 12: 266–73. 34. Mackey DA, Mackinnon JR, Brown SA et al. Twins eye
27. Seddon JM, Sahagian CR, Glynn RJ, Sperduto RD, study in Tasmania (TEST): rationale and methodology
Gragoudas ES. Evaluation of an iris color classification to recruit and examine twins. Twin Res Hum Genet 2009;
system. The Eye Disorders Case-Control Study Group. 12: 441–54.
Invest Ophthalmol Vis Sci 1990; 31: 1592–8. 35. Macdonald W, Newnham J, Gurrin L, Evans S. Effect
28. Takamoto T, Schwartz B, Cantor LB, Hoop JS, Steffens of frequent prenatal ultrasound on birthweight: follow
T. Measurement of iris color using computerized image up at 1 year of age. Western Australian Pregnancy
analysis. Curr Eye Res 2001; 22: 412–19. Cohort (Raine) Working Group. Lancet 1996; 348: 482.

© 2011 The Authors


Clinical and Experimental Ophthalmology © 2011 Royal Australian and New Zealand College of Ophthalmologists

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