Abstract

We measured the amplitude of the directional component of the bleached fundus reflectance, the so-called optical Stiles–Crawford effect, as a function of wavelength. The directional reflectance originates from within the outer segments of the photoreceptors. Thus only two anterior absorbers are of importance: macular pigment and the crystalline lens. Analysis of spectra obtained in pseudophakes established that the cone photoreceptors act as spectrally neutral reflectors. The reflectance spectra, expressed in density units, resembled the macular pigment density spectrum. Studying age effects in the lens of normal subjects resulted in a description of the optical density of the lens in terms of a “young” and an “aged” template. The young template represents the pigment O-β-glucoside of 3-hydroxykynurenine, which dominates the light absorption in young eyes and decreases with age. The aged template represents the pigments accumulating in the lens with age. The total optical density increased with age, but it was lower in the wavelength region 500–650 nm than was previously assumed on the basis of psychophysical studies. Analysis of the spectra also provided precise individual estimates of the optical density of macular pigment. Finally, we observed a decrease in the photoreceptor reflectivity with age, possibly reflecting a degradation of the photoreceptors.

© 2004 Optical Society of America

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2003 (2)

T. T. J. M. Berendschot, P. J. DeLint, D. van Norren, “Fundus reflectance—historical and present ideas,” Prog. Retin. Eye Res. 22, 171–200 (2003).
[CrossRef] [PubMed]

N. P. A. Zagers, T. T. J. M. Berendschot, D. van Norren, “Wavelength dependence of reflectometric cone photoreceptor directionality,” J. Opt. Soc. Am. A 20, 18–23 (2003).
[CrossRef]

2002 (5)

N. P. A. Zagers, J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “Simultaneous measurement of foveal spectral reflectance and cone-photoreceptor directionality,” Appl. Opt. 41, 4686–4696 (2002).
[CrossRef] [PubMed]

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

A. Roorda, D. R. Williams, “Optical fiber properties of individual human cones,” J. Vision 2, 404–412 (2002).
[CrossRef]

M. Hammer, D. Schweitzer, “Quantitative reflection spectroscopy at the human ocular fundus,” Phys. Med. Biol. 47, 179–191 (2002).
[CrossRef] [PubMed]

S. J. Preece, E. Claridge, “Monte Carlo modelling of the spectral reflectance of the human eye,” Phys. Med. Biol. 47, 2863–2877 (2002).
[CrossRef] [PubMed]

2001 (6)

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

R. C. Heckathorn, J. Dillon, E. R. Gaillard, “Synthesis and purification of 3-hydroxykynurenine-O-β-glucoside, a primate lens ultraviolet filter, and its application in a two-step assay for β-glucosidase activity,” Anal. Biochem. 299, 78–83 (2001).
[CrossRef] [PubMed]

G. L. Savage, C. A. Johnson, D. L. Howard, “A comparison of noninvasive objective and subjective measurements of the optical density of human ocular media,” Optom. Vision Sci. 78, 386–395 (2001).
[CrossRef]

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

F. C. Delori, D. G. Goger, B. R. Hammond, D. M. Snodderly, S. A. Burns, “Macular pigment density measured by autofluorescence spectrometry: comparison with reflectometry and heterochromatic flicker photometry,” J. Opt. Soc. Am. A 18, 1212–1230 (2001).
[CrossRef]

2000 (2)

B. R. Hammond, M. Caruso-Avery, “Macular pigment optical density in a Southwestern sample,” Invest. Ophthalmol. Visual Sci. 41, 1492–1497 (2000).

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

1999 (4)

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
[CrossRef] [PubMed]

A. Stockman, L. T. Sharpe, C. Fach, “The spectral sensitivity of the human short-wavelength sensitive cones derived from thresholds and color matches,” Vision Res. 39, 2901–2927 (1999).
[CrossRef] [PubMed]

S. Marcos, S. A. Burns, “Cone spacing and waveguide properties from cone directionality measurements,” J. Opt. Soc. Am. A 16, 995–1004 (1999).
[CrossRef]

J. C. He, S. Marcos, S. A. Burns, “Comparison of cone directionality determined by psychophysical and reflectometric techniques,” J. Opt. Soc. Am. A 16, 2363–2369 (1999).
[CrossRef]

1998 (1)

1997 (3)

J. Xu, J. Pokorny, V. C. Smith, “Optical density of the human lens,” J. Opt. Soc. Am. A 14, 953–960 (1997).
[CrossRef]

P. J. DeLint, T. T. J. M. Berendschot, D. van Norren, “Local photoreceptor alignment measured with a scanning laser ophthalmoscope,” Vision Res. 37, 243–248 (1997).
[CrossRef] [PubMed]

J. M. Gorrand, F. C. Delori, “A model for assessment of cone directionality,” J. Mod. Opt. 44, 473–491 (1997).
[CrossRef]

1996 (2)

J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “The pathways of light measured in fundus reflectometry,” Vision Res. 36, 2229–2247 (1996).
[CrossRef] [PubMed]

F. C. Delori, S. A. Burns, “Fundus reflectance and the measurement of crystalline lens density,” J. Opt. Soc. Am. A 13, 215–226 (1996).
[CrossRef]

1995 (4)

S. A. Burns, S. Wu, F. Delori, A. E. Elsner, “Direct measurement of human-cone-photoreceptor alignment,” J. Opt. Soc. Am. A 12, 2329–2338 (1995).
[CrossRef]

T. J. T. P. van den Berg, J. K. IJspeert, “Light scattering in donor lenses,” Vision Res. 35, 169–177 (1995).
[CrossRef] [PubMed]

J.-M. Gorrand, F. Delori, “A reflectometric technique for assessing photoreceptor alignment,” Vision Res. 35, 999–1010 (1995).
[CrossRef] [PubMed]

T. J. T. P. van den Berg, J. Felius, “Relationship between spectral transmittance and slit lamp color of human lenses,” Invest. Ophthalmol. Visual Sci. 36, 322–329 (1995).

1994 (1)

T. J. T. P. van den Berg, K. E. W. P. Tan, “Light transmittance of the human cornea from 320 to 700 nm for different ages,” Vision Res. 34, 1453–1456 (1994).
[CrossRef] [PubMed]

1993 (7)

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
[CrossRef]

G. M. Stutchbury, R. J. W. Truscott, “The modification of proteins by 3-hydroxykynurenine,” Exp. Eye Res. 57, 149–155 (1993).
[CrossRef] [PubMed]

D. Whitaker, R. Steen, D. B. Elliott, “Light scatter in the normal young, elderly, and cataractous eye demonstrates little wavelength dependency,” Optom. Vision Sci. 70, 963–968 (1993).
[CrossRef]

M. J. Piket-May, A. Taflove, J. B. Troy, “Electrodynamics of visible-light interactions with the vertebrate retinal rod,” Opt. Lett. 18, 568–570 (1993).
[CrossRef] [PubMed]

A. M. Wood, R. J. W. Truscott, “UV filters in human lenses: tryptophan catabolism,” Exp. Eye Res. 56, 317–325 (1993).
[CrossRef] [PubMed]

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

1991 (2)

J. Dillon, “The photophysics and photobiology of the eye,” J. Photochem. Photobiol. B 10, 23–40 (1991).
[CrossRef] [PubMed]

R. A. Weale, “The lenticular nucleus, light, and the retina,” Exp. Eye Res. 53, 213–218 (1991).
[CrossRef] [PubMed]

1990 (1)

J. Dillon, R.-H. Wang, S. J. Atherton, “Photochemical and photophysical studies on human lens constituents,” Photochem. Photobiol. 52, 849–854 (1990).
[CrossRef] [PubMed]

1989 (1)

1988 (3)

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

R. A. Weale, “Age and the transmittance of the human crystalline lens,” J. Physiol. (London) 395, 577–587 (1988).

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

1987 (4)

J. S. Werner, S. K. Donnelly, R. Kliegl, “Aging and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering,” Vision Res. 27, 257–268 (1987).
[CrossRef] [PubMed]

B. R. Wooten, G. A. Geri, “Psychophysical determination of intraocular light scatter as a function of wavelength,” Vision Res. 27, 1291–1298 (1987).
[CrossRef] [PubMed]

Y. Murata, “Light absorption characteristics of the lens capsule,” Ophthalmic Res. 19, 107–112 (1987).
[CrossRef] [PubMed]

J. Pokorny, V. C. Smith, M. Lutze, “Aging of the human lens,” Appl. Opt. 26, 1437–1440 (1987).
[CrossRef] [PubMed]

1986 (3)

D. van Norren, L. F. Tiemeijer, “Spectral reflectance of the human eye,” Vision Res. 26, 313–320 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, D. van Norren, “Intensity and polarization of light scattered at small angles from the human fovea,” Vision Res. 26, 485–494 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, “Directionality and alignment of the foveal receptors, assessed with light scattered from the human fundus in vivo,” Vision Res. 26, 495–500 (1986).
[CrossRef] [PubMed]

1984 (3)

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

D. M. Snodderly, J. D. Auran, F. C. Delori, “The macular pigment. II. Spatial distribution in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 674–685 (1984).

J. Dillon, “Photolytic changes in lens proteins,” Curr. Eye Res. 3, 145–150 (1984).
[CrossRef] [PubMed]

1981 (1)

M. Bando, A. Nakajima, K. Satoh, “Spectrophotometric estimation of 3-OH L-kynurenine O-β-glucoside in the human lens,” J. Biochem. 89, 103–109 (1981).
[PubMed]

1975 (1)

J. J. Vos, F. L. van Os, “The effect of lens density on the Stiles–Crawford effect,” Vision Res. 15, 749–751 (1975).
[CrossRef] [PubMed]

1974 (1)

D. van Norren, J. J. Vos, “Spectral transmission of the human ocular media,” Vision Res. 14, 1237–1244 (1974).
[CrossRef]

1972 (1)

S. Coren, J. S. Girgus, “Density of human lens pigmentation: in vivo measures over an extended age range,” Vision Res. 12, 343–346 (1972).
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1971 (3)

S. Zigman, “Eye lens color: formation and function,” Science 171, 807–809 (1971).
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J. Mellerio, “Light absorption and scatter in the human lens,” Vision Res. 11, 129–141 (1971).
[CrossRef] [PubMed]

R. van Heyningen, “Fluorescent glucoside in the human lens,” Nature (London) 230, 393–394 (1971).
[CrossRef]

1969 (1)

G. F. Cooper, J. G. Robson, “The yellow colour of the lens of man and other primates,” J. Physiol. (London) 203, 411–417 (1969).

1962 (1)

E. A. Boettner, J. R. Wolter, “Transmission of the ocular media,” Invest. Ophthalmol. 1, 776–783 (1962).

1959 (1)

F. S. Said, R. A. Weale, “The variation with age of the spectral transmissivity of the living human crystalline lens,” Gerontologia (Basel) 3, 213–131 (1959).
[CrossRef]

Adams, A. J.

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

Allen, K. A.

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

Atherton, S. J.

J. Dillon, R.-H. Wang, S. J. Atherton, “Photochemical and photophysical studies on human lens constituents,” Photochem. Photobiol. 52, 849–854 (1990).
[CrossRef] [PubMed]

Auran, J. D.

D. M. Snodderly, J. D. Auran, F. C. Delori, “The macular pigment. II. Spatial distribution in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 674–685 (1984).

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

Bando, M.

M. Bando, A. Nakajima, K. Satoh, “Spectrophotometric estimation of 3-OH L-kynurenine O-β-glucoside in the human lens,” J. Biochem. 89, 103–109 (1981).
[PubMed]

Beatty, S.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Berendschot, T. T. J. M.

N. P. A. Zagers, T. T. J. M. Berendschot, D. van Norren, “Wavelength dependence of reflectometric cone photoreceptor directionality,” J. Opt. Soc. Am. A 20, 18–23 (2003).
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T. T. J. M. Berendschot, P. J. DeLint, D. van Norren, “Fundus reflectance—historical and present ideas,” Prog. Retin. Eye Res. 22, 171–200 (2003).
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N. P. A. Zagers, J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “Simultaneous measurement of foveal spectral reflectance and cone-photoreceptor directionality,” Appl. Opt. 41, 4686–4696 (2002).
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W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

P. J. DeLint, T. T. J. M. Berendschot, D. van Norren, “Local photoreceptor alignment measured with a scanning laser ophthalmoscope,” Vision Res. 37, 243–248 (1997).
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J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “The pathways of light measured in fundus reflectometry,” Vision Res. 36, 2229–2247 (1996).
[CrossRef] [PubMed]

Boettner, E. A.

E. A. Boettner, J. R. Wolter, “Transmission of the ocular media,” Invest. Ophthalmol. 1, 776–783 (1962).

E. A. Boettner, Spectral Transmission of the Eye [Report of the University of Michigan, contract AF41(609)-2966, Ann Arbor, Mich., 1967].

Bone, R. A.

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

Boulton, M. E.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Bova, L. M.

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

Boynton, R. M.

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

Broekmans, W. M. R.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Brown, P. K.

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

Burns, S. A.

Carden, D.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Caruso-Avery, M.

B. R. Hammond, M. Caruso-Avery, “Macular pigment optical density in a Southwestern sample,” Invest. Ophthalmol. Visual Sci. 41, 1492–1497 (2000).

Ciulla, T. A.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Claridge, E.

S. J. Preece, E. Claridge, “Monte Carlo modelling of the spectral reflectance of the human eye,” Phys. Med. Biol. 47, 2863–2877 (2002).
[CrossRef] [PubMed]

Cooper, D. A.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Cooper, G. F.

G. F. Cooper, J. G. Robson, “The yellow colour of the lens of man and other primates,” J. Physiol. (London) 203, 411–417 (1969).

Coren, S.

S. Coren, J. S. Girgus, “Density of human lens pigmentation: in vivo measures over an extended age range,” Vision Res. 12, 343–346 (1972).
[CrossRef] [PubMed]

Curcio, C. A.

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

Curran-Celantano, J.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Danis, R. P.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
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de Vries, A. J.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

DeLint, P. J.

T. T. J. M. Berendschot, P. J. DeLint, D. van Norren, “Fundus reflectance—historical and present ideas,” Prog. Retin. Eye Res. 22, 171–200 (2003).
[CrossRef] [PubMed]

P. J. DeLint, T. T. J. M. Berendschot, D. van Norren, “Local photoreceptor alignment measured with a scanning laser ophthalmoscope,” Vision Res. 37, 243–248 (1997).
[CrossRef] [PubMed]

Delori, F.

J.-M. Gorrand, F. Delori, “A reflectometric technique for assessing photoreceptor alignment,” Vision Res. 35, 999–1010 (1995).
[CrossRef] [PubMed]

S. A. Burns, S. Wu, F. Delori, A. E. Elsner, “Direct measurement of human-cone-photoreceptor alignment,” J. Opt. Soc. Am. A 12, 2329–2338 (1995).
[CrossRef]

Delori, F. C.

F. C. Delori, D. G. Goger, B. R. Hammond, D. M. Snodderly, S. A. Burns, “Macular pigment density measured by autofluorescence spectrometry: comparison with reflectometry and heterochromatic flicker photometry,” J. Opt. Soc. Am. A 18, 1212–1230 (2001).
[CrossRef]

J. M. Gorrand, F. C. Delori, “A model for assessment of cone directionality,” J. Mod. Opt. 44, 473–491 (1997).
[CrossRef]

F. C. Delori, S. A. Burns, “Fundus reflectance and the measurement of crystalline lens density,” J. Opt. Soc. Am. A 13, 215–226 (1996).
[CrossRef]

F. C. Delori, K. P. Pflibsen, “Spectral reflectance of the human ocular fundus,” Appl. Opt. 28, 1061–1077 (1989).
[CrossRef] [PubMed]

D. M. Snodderly, J. D. Auran, F. C. Delori, “The macular pigment. II. Spatial distribution in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 674–685 (1984).

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

S. A. Burns, J. C. He, F. C. Delori, “Do the cones see light scattered from the deeper retinal layers?” in Vision Science and Its Applications, Vol. 1 of 1997 OSA Technical Digest Series (Optical Society of America, Washington D.C., 1997), pp. 94–97.

Dillon, J.

R. C. Heckathorn, J. Dillon, E. R. Gaillard, “Synthesis and purification of 3-hydroxykynurenine-O-β-glucoside, a primate lens ultraviolet filter, and its application in a two-step assay for β-glucosidase activity,” Anal. Biochem. 299, 78–83 (2001).
[CrossRef] [PubMed]

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
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J. Dillon, “The photophysics and photobiology of the eye,” J. Photochem. Photobiol. B 10, 23–40 (1991).
[CrossRef] [PubMed]

J. Dillon, R.-H. Wang, S. J. Atherton, “Photochemical and photophysical studies on human lens constituents,” Photochem. Photobiol. 52, 849–854 (1990).
[CrossRef] [PubMed]

J. Dillon, “Photolytic changes in lens proteins,” Curr. Eye Res. 3, 145–150 (1984).
[CrossRef] [PubMed]

Donnelly, S. K.

J. S. Werner, S. K. Donnelly, R. Kliegl, “Aging and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering,” Vision Res. 27, 257–268 (1987).
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Elliott, D. B.

D. Whitaker, R. Steen, D. B. Elliott, “Light scatter in the normal young, elderly, and cataractous eye demonstrates little wavelength dependency,” Optom. Vision Sci. 70, 963–968 (1993).
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Elsner, A. E.

Esterson, F. D.

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

Fach, C.

A. Stockman, L. T. Sharpe, C. Fach, “The spectral sensitivity of the human short-wavelength sensitive cones derived from thresholds and color matches,” Vision Res. 39, 2901–2927 (1999).
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Felius, J.

T. J. T. P. van den Berg, J. Felius, “Relationship between spectral transmittance and slit lamp color of human lenses,” Invest. Ophthalmol. Visual Sci. 36, 322–329 (1995).

Fernandez, L.

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

Filloon, T. G.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Flannery, B. P.

W. H. Press, B. P. Flannery, S. A. Teukolsky, W. T. Vetterling, Numerical Recipes in Pascal (Cambridge U. Press, Cambridge, UK, 1989).

Gaillard, E. R.

R. C. Heckathorn, J. Dillon, E. R. Gaillard, “Synthesis and purification of 3-hydroxykynurenine-O-β-glucoside, a primate lens ultraviolet filter, and its application in a two-step assay for β-glucosidase activity,” Anal. Biochem. 299, 78–83 (2001).
[CrossRef] [PubMed]

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
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Geri, G. A.

B. R. Wooten, G. A. Geri, “Psychophysical determination of intraocular light scatter as a function of wavelength,” Vision Res. 27, 1291–1298 (1987).
[CrossRef] [PubMed]

Girgus, J. S.

S. Coren, J. S. Girgus, “Density of human lens pigmentation: in vivo measures over an extended age range,” Vision Res. 12, 343–346 (1972).
[CrossRef] [PubMed]

Goger, D. G.

Goldbohm, R. A.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Gorrand, J. M.

J. M. Gorrand, F. C. Delori, “A model for assessment of cone directionality,” J. Mod. Opt. 44, 473–491 (1997).
[CrossRef]

Gorrand, J.-M.

J.-M. Gorrand, F. Delori, “A reflectometric technique for assessing photoreceptor alignment,” Vision Res. 35, 999–1010 (1995).
[CrossRef] [PubMed]

Haegerstrom-Portnoy, G.

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

Hammer, M.

M. Hammer, D. Schweitzer, “Quantitative reflection spectroscopy at the human ocular fundus,” Phys. Med. Biol. 47, 179–191 (2002).
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Hammond, B. R.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

F. C. Delori, D. G. Goger, B. R. Hammond, D. M. Snodderly, S. A. Burns, “Macular pigment density measured by autofluorescence spectrometry: comparison with reflectometry and heterochromatic flicker photometry,” J. Opt. Soc. Am. A 18, 1212–1230 (2001).
[CrossRef]

B. R. Hammond, M. Caruso-Avery, “Macular pigment optical density in a Southwestern sample,” Invest. Ophthalmol. Visual Sci. 41, 1492–1497 (2000).

He, J. C.

J. C. He, S. Marcos, S. A. Burns, “Comparison of cone directionality determined by psychophysical and reflectometric techniques,” J. Opt. Soc. Am. A 16, 2363–2369 (1999).
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S. Marcos, S. A. Burns, J. C. He, “Model for cone directionality reflectometric measurements based on scattering,” J. Opt. Soc. Am. A 15, 2012–2022 (1998).
[CrossRef]

S. A. Burns, J. C. He, F. C. Delori, “Do the cones see light scattered from the deeper retinal layers?” in Vision Science and Its Applications, Vol. 1 of 1997 OSA Technical Digest Series (Optical Society of America, Washington D.C., 1997), pp. 94–97.

Heckathorn, R. C.

R. C. Heckathorn, J. Dillon, E. R. Gaillard, “Synthesis and purification of 3-hydroxykynurenine-O-β-glucoside, a primate lens ultraviolet filter, and its application in a two-step assay for β-glucosidase activity,” Anal. Biochem. 299, 78–83 (2001).
[CrossRef] [PubMed]

Henson, D. B.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Hewlett, S. E.

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

Howard, D. L.

G. L. Savage, C. A. Johnson, D. L. Howard, “A comparison of noninvasive objective and subjective measurements of the optical density of human ocular media,” Optom. Vision Sci. 78, 386–395 (2001).
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C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
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IJspeert, J. K.

T. J. T. P. van den Berg, J. K. IJspeert, “Light scattering in donor lenses,” Vision Res. 35, 169–177 (1995).
[CrossRef] [PubMed]

Jamie, J. F.

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

Johnson, C. A.

G. L. Savage, C. A. Johnson, D. L. Howard, “A comparison of noninvasive objective and subjective measurements of the optical density of human ocular media,” Optom. Vision Sci. 78, 386–395 (2001).
[CrossRef]

C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
[CrossRef]

Kalina, R. E.

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

Kardinaal, A. F. M.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Kliegl, R.

J. S. Werner, S. K. Donnelly, R. Kliegl, “Aging and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering,” Vision Res. 27, 257–268 (1987).
[CrossRef] [PubMed]

Klöpping-Ketelaars, I. A. A.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Koh, H.-H.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Krauskopf, J.

J. Krauskopf, “Some experiments with a photoelectric ophthalmoscope,” in Performance of the Eye at Low Luminances, M. A. Bouman, J. J. Vos, eds. (Excerpta Medica, Amsterdam, 1966), pp. 171–181.

Landrum, J. T.

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

Lutze, M.

Marcos, S.

Marshall, D.

C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
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Marshall, J.

J. Marshall, “Ageing changes in human cones,” in International Congress Series, K. Shimizu, J. A. Oosterhuis, eds. (Excerpta Media, Amsterdam, 1979), pp. 375–378.

Mellerio, J.

J. Mellerio, “Light absorption and scatter in the human lens,” Vision Res. 11, 129–141 (1971).
[CrossRef] [PubMed]

Merriam, J. C.

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
[CrossRef] [PubMed]

Millican, C. L.

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

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Y. Murata, “Light absorption characteristics of the lens capsule,” Ophthalmic Res. 19, 107–112 (1987).
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Murray, I. J.

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

Nakajima, A.

M. Bando, A. Nakajima, K. Satoh, “Spectrophotometric estimation of 3-OH L-kynurenine O-β-glucoside in the human lens,” J. Biochem. 89, 103–109 (1981).
[PubMed]

Pflibsen, K. P.

Piket-May, M. J.

Pokorny, J.

Pratt, L. M.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Preece, S. J.

S. J. Preece, E. Claridge, “Monte Carlo modelling of the spectral reflectance of the human eye,” Phys. Med. Biol. 47, 2863–2877 (2002).
[CrossRef] [PubMed]

Press, W. H.

W. H. Press, B. P. Flannery, S. A. Teukolsky, W. T. Vetterling, Numerical Recipes in Pascal (Cambridge U. Press, Cambridge, UK, 1989).

Riccardi, K. A.

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Robson, J. G.

G. F. Cooper, J. G. Robson, “The yellow colour of the lens of man and other primates,” J. Physiol. (London) 203, 411–417 (1969).

Roorda, A.

A. Roorda, D. R. Williams, “Optical fiber properties of individual human cones,” J. Vision 2, 404–412 (2002).
[CrossRef]

Said, F. S.

F. S. Said, R. A. Weale, “The variation with age of the spectral transmissivity of the living human crystalline lens,” Gerontologia (Basel) 3, 213–131 (1959).
[CrossRef]

Sample, P. A.

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

Satoh, K.

M. Bando, A. Nakajima, K. Satoh, “Spectrophotometric estimation of 3-OH L-kynurenine O-β-glucoside in the human lens,” J. Biochem. 89, 103–109 (1981).
[PubMed]

Savage, G. L.

G. L. Savage, C. A. Johnson, D. L. Howard, “A comparison of noninvasive objective and subjective measurements of the optical density of human ocular media,” Optom. Vision Sci. 78, 386–395 (2001).
[CrossRef]

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

Schweitzer, D.

M. Hammer, D. Schweitzer, “Quantitative reflection spectroscopy at the human ocular fundus,” Phys. Med. Biol. 47, 179–191 (2002).
[CrossRef] [PubMed]

Sharpe, L. T.

A. Stockman, L. T. Sharpe, C. Fach, “The spectral sensitivity of the human short-wavelength sensitive cones derived from thresholds and color matches,” Vision Res. 39, 2901–2927 (1999).
[CrossRef] [PubMed]

Shu, H.

C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
[CrossRef]

Smith, V. C.

Snodderly, D. M.

F. C. Delori, D. G. Goger, B. R. Hammond, D. M. Snodderly, S. A. Burns, “Macular pigment density measured by autofluorescence spectrometry: comparison with reflectometry and heterochromatic flicker photometry,” J. Opt. Soc. Am. A 18, 1212–1230 (2001).
[CrossRef]

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

D. M. Snodderly, J. D. Auran, F. C. Delori, “The macular pigment. II. Spatial distribution in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 674–685 (1984).

Steen, R.

D. Whitaker, R. Steen, D. B. Elliott, “Light scatter in the normal young, elderly, and cataractous eye demonstrates little wavelength dependency,” Optom. Vision Sci. 70, 963–968 (1993).
[CrossRef]

Stiles, W. S.

G. Wyszecki, W. S. Stiles, Color Science: Concepts and Methods, Quantitative Data and Formulae, 2nd ed. (Wiley, New York, 1982).

Stockman, A.

A. Stockman, L. T. Sharpe, C. Fach, “The spectral sensitivity of the human short-wavelength sensitive cones derived from thresholds and color matches,” Vision Res. 39, 2901–2927 (1999).
[CrossRef] [PubMed]

Stutchbury, G. M.

G. M. Stutchbury, R. J. W. Truscott, “The modification of proteins by 3-hydroxykynurenine,” Exp. Eye Res. 57, 149–155 (1993).
[CrossRef] [PubMed]

Sweeney, M. H. J.

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

Taflove, A.

Tan, K. E. W. P.

T. J. T. P. van den Berg, K. E. W. P. Tan, “Light transmittance of the human cornea from 320 to 700 nm for different ages,” Vision Res. 34, 1453–1456 (1994).
[CrossRef] [PubMed]

K. E. W. P. Tan, “Vision in the ultraviolet,” Ph.D thesis (Universiteit Utrecht, Utrecht, The Netherlands, 1971).

Tarsis, S. L.

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

Teukolsky, S. A.

W. H. Press, B. P. Flannery, S. A. Teukolsky, W. T. Vetterling, Numerical Recipes in Pascal (Cambridge U. Press, Cambridge, UK, 1989).

Tiemeijer, L. F.

D. van Norren, L. F. Tiemeijer, “Spectral reflectance of the human eye,” Vision Res. 26, 313–320 (1986).
[CrossRef] [PubMed]

Tijburg, L. B. M.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Troy, J. B.

Truscott, R. J. W.

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

G. M. Stutchbury, R. J. W. Truscott, “The modification of proteins by 3-hydroxykynurenine,” Exp. Eye Res. 57, 149–155 (1993).
[CrossRef] [PubMed]

A. M. Wood, R. J. W. Truscott, “UV filters in human lenses: tryptophan catabolism,” Exp. Eye Res. 56, 317–325 (1993).
[CrossRef] [PubMed]

van Blokland, G. J.

G. J. van Blokland, D. van Norren, “Intensity and polarization of light scattered at small angles from the human fovea,” Vision Res. 26, 485–494 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, “Directionality and alignment of the foveal receptors, assessed with light scattered from the human fundus in vivo,” Vision Res. 26, 495–500 (1986).
[CrossRef] [PubMed]

van de Kraats, J.

van den Berg, T. J. T. P.

T. J. T. P. van den Berg, J. K. IJspeert, “Light scattering in donor lenses,” Vision Res. 35, 169–177 (1995).
[CrossRef] [PubMed]

T. J. T. P. van den Berg, J. Felius, “Relationship between spectral transmittance and slit lamp color of human lenses,” Invest. Ophthalmol. Visual Sci. 36, 322–329 (1995).

T. J. T. P. van den Berg, K. E. W. P. Tan, “Light transmittance of the human cornea from 320 to 700 nm for different ages,” Vision Res. 34, 1453–1456 (1994).
[CrossRef] [PubMed]

van Heyningen, R.

R. van Heyningen, “Fluorescent glucoside in the human lens,” Nature (London) 230, 393–394 (1971).
[CrossRef]

van Norren, D.

T. T. J. M. Berendschot, P. J. DeLint, D. van Norren, “Fundus reflectance—historical and present ideas,” Prog. Retin. Eye Res. 22, 171–200 (2003).
[CrossRef] [PubMed]

N. P. A. Zagers, T. T. J. M. Berendschot, D. van Norren, “Wavelength dependence of reflectometric cone photoreceptor directionality,” J. Opt. Soc. Am. A 20, 18–23 (2003).
[CrossRef]

N. P. A. Zagers, J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “Simultaneous measurement of foveal spectral reflectance and cone-photoreceptor directionality,” Appl. Opt. 41, 4686–4696 (2002).
[CrossRef] [PubMed]

P. J. DeLint, T. T. J. M. Berendschot, D. van Norren, “Local photoreceptor alignment measured with a scanning laser ophthalmoscope,” Vision Res. 37, 243–248 (1997).
[CrossRef] [PubMed]

J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “The pathways of light measured in fundus reflectometry,” Vision Res. 36, 2229–2247 (1996).
[CrossRef] [PubMed]

D. van Norren, L. F. Tiemeijer, “Spectral reflectance of the human eye,” Vision Res. 26, 313–320 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, D. van Norren, “Intensity and polarization of light scattered at small angles from the human fovea,” Vision Res. 26, 485–494 (1986).
[CrossRef] [PubMed]

D. van Norren, J. J. Vos, “Spectral transmission of the human ocular media,” Vision Res. 14, 1237–1244 (1974).
[CrossRef]

van Os, F. L.

J. J. Vos, F. L. van Os, “The effect of lens density on the Stiles–Crawford effect,” Vision Res. 15, 749–751 (1975).
[CrossRef] [PubMed]

van Poppel, G.

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Vetterling, W. T.

W. H. Press, B. P. Flannery, S. A. Teukolsky, W. T. Vetterling, Numerical Recipes in Pascal (Cambridge U. Press, Cambridge, UK, 1989).

Vos, J. J.

J. J. Vos, F. L. van Os, “The effect of lens density on the Stiles–Crawford effect,” Vision Res. 15, 749–751 (1975).
[CrossRef] [PubMed]

D. van Norren, J. J. Vos, “Spectral transmission of the human ocular media,” Vision Res. 14, 1237–1244 (1974).
[CrossRef]

J. J. Vos, Literature Review of Human Macular Absorption in the Visible and Its Consequences for the Cone Receptor Primaries (National Organization for Applied Scientific Research in The Netherlands, Institute for Perception RVO-TNO, Soesterberg, The Netherlands, 1972).

Walraven, P. L.

P. L. Walraven, , Draft 7 (personal communication, 2003) (plwalraven@hotmail.com).

Wang, R.-H.

J. Dillon, R.-H. Wang, S. J. Atherton, “Photochemical and photophysical studies on human lens constituents,” Photochem. Photobiol. 52, 849–854 (1990).
[CrossRef] [PubMed]

Weale, R. A.

R. A. Weale, “The lenticular nucleus, light, and the retina,” Exp. Eye Res. 53, 213–218 (1991).
[CrossRef] [PubMed]

R. A. Weale, “Age and the transmittance of the human crystalline lens,” J. Physiol. (London) 395, 577–587 (1988).

F. S. Said, R. A. Weale, “The variation with age of the spectral transmissivity of the living human crystalline lens,” Gerontologia (Basel) 3, 213–131 (1959).
[CrossRef]

Weinreb, R. N.

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

Werner, J. S.

J. S. Werner, S. K. Donnelly, R. Kliegl, “Aging and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering,” Vision Res. 27, 257–268 (1987).
[CrossRef] [PubMed]

Whitaker, D.

D. Whitaker, R. Steen, D. B. Elliott, “Light scatter in the normal young, elderly, and cataractous eye demonstrates little wavelength dependency,” Optom. Vision Sci. 70, 963–968 (1993).
[CrossRef]

Williams, D. R.

A. Roorda, D. R. Williams, “Optical fiber properties of individual human cones,” J. Vision 2, 404–412 (2002).
[CrossRef]

Wolter, J. R.

E. A. Boettner, J. R. Wolter, “Transmission of the ocular media,” Invest. Ophthalmol. 1, 776–783 (1962).

Wood, A. M.

A. M. Wood, R. J. W. Truscott, “UV filters in human lenses: tryptophan catabolism,” Exp. Eye Res. 56, 317–325 (1993).
[CrossRef] [PubMed]

Wooten, B. R.

B. R. Wooten, G. A. Geri, “Psychophysical determination of intraocular light scatter as a function of wavelength,” Vision Res. 27, 1291–1298 (1987).
[CrossRef] [PubMed]

Wu, S.

Wyszecki, G.

G. Wyszecki, W. S. Stiles, Color Science: Concepts and Methods, Quantitative Data and Formulae, 2nd ed. (Wiley, New York, 1982).

Xu, J.

Zagers, N. P. A.

Zheng, L.

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
[CrossRef] [PubMed]

Zigman, S.

S. Zigman, “Eye lens color: formation and function,” Science 171, 807–809 (1971).
[CrossRef] [PubMed]

Am. J. Clin. Nutr. (1)

W. M. R. Broekmans, T. T. J. M. Berendschot, I. A. A. Klöpping-Ketelaars, A. J. de Vries, R. A. Goldbohm, L. B. M. Tijburg, A. F. M. Kardinaal, G. van Poppel, “Macular pigment density in relation to serum and adipose tissue concentrations of lutein and serum concentrations of zeaxanthin,” Am. J. Clin. Nutr. 76, 595–603 (2002).
[PubMed]

Anal. Biochem. (1)

R. C. Heckathorn, J. Dillon, E. R. Gaillard, “Synthesis and purification of 3-hydroxykynurenine-O-β-glucoside, a primate lens ultraviolet filter, and its application in a two-step assay for β-glucosidase activity,” Anal. Biochem. 299, 78–83 (2001).
[CrossRef] [PubMed]

Appl. Opt. (3)

Clin. Vision Sci. (1)

G. L. Savage, G. Haegerstrom-Portnoy, A. J. Adams, S. E. Hewlett, “Age changes in the optical density of human ocular media,” Clin. Vision Sci. 8, 97–108 (1993).

Curr. Eye Res. (1)

J. Dillon, “Photolytic changes in lens proteins,” Curr. Eye Res. 3, 145–150 (1984).
[CrossRef] [PubMed]

Exp. Eye Res. (4)

R. A. Weale, “The lenticular nucleus, light, and the retina,” Exp. Eye Res. 53, 213–218 (1991).
[CrossRef] [PubMed]

A. M. Wood, R. J. W. Truscott, “UV filters in human lenses: tryptophan catabolism,” Exp. Eye Res. 56, 317–325 (1993).
[CrossRef] [PubMed]

G. M. Stutchbury, R. J. W. Truscott, “The modification of proteins by 3-hydroxykynurenine,” Exp. Eye Res. 57, 149–155 (1993).
[CrossRef] [PubMed]

J. Dillon, L. Zheng, J. C. Merriam, E. R. Gaillard, “The optical properties of the anterior segment of the eye: implications for cortical cataract,” Exp. Eye Res. 68, 785–795 (1999).
[CrossRef] [PubMed]

Gerontologia (Basel) (1)

F. S. Said, R. A. Weale, “The variation with age of the spectral transmissivity of the living human crystalline lens,” Gerontologia (Basel) 3, 213–131 (1959).
[CrossRef]

Invest. Ophthalmol. (1)

E. A. Boettner, J. R. Wolter, “Transmission of the ocular media,” Invest. Ophthalmol. 1, 776–783 (1962).

Invest. Ophthalmol. Visual Sci. (10)

C. A. Curcio, C. L. Millican, K. A. Allen, R. E. Kalina, “Aging of the human photoreceptor mosaic: evidence for selective vulnerability of rods in central retina,” Invest. Ophthalmol. Visual Sci. 34, 3278–3296 (1993).

B. R. Hammond, M. Caruso-Avery, “Macular pigment optical density in a Southwestern sample,” Invest. Ophthalmol. Visual Sci. 41, 1492–1497 (2000).

S. Beatty, I. J. Murray, D. B. Henson, D. Carden, H.-H. Koh, M. E. Boulton, “Macular pigment and risk for age-related macular degeneration in subjects from a Northern European population,” Invest. Ophthalmol. Visual Sci. 42, 439–446 (2001).

P. A. Sample, F. D. Esterson, R. N. Weinreb, R. M. Boynton, “The aging lens: in vivo assessment of light absorption in 84 human eyes,” Invest. Ophthalmol. Visual Sci. 29, 1306–1311 (1988).

R. A. Bone, J. T. Landrum, L. Fernandez, S. L. Tarsis, “Analysis of the macular pigment by HPLC: retinal distribution and age study,” Invest. Ophthalmol. Visual Sci. 29, 843–849 (1988).

E. R. Gaillard, L. Zheng, J. C. Merriam, J. Dillon, “Age-related changes in the absorption characteristics of the primate lens,” Invest. Ophthalmol. Visual Sci. 41, 1454–1459 (2000).

L. M. Bova, M. H. J. Sweeney, J. F. Jamie, R. J. W. Truscott, “Major changes in human ocular UV protection withage,” Invest. Ophthalmol. Visual Sci. 42, 200–205 (2001).

T. J. T. P. van den Berg, J. Felius, “Relationship between spectral transmittance and slit lamp color of human lenses,” Invest. Ophthalmol. Visual Sci. 36, 322–329 (1995).

D. M. Snodderly, P. K. Brown, F. C. Delori, J. D. Auran, “The macular pigment. I. Absorbance spectra, localization, and discrimination from other yellow pigments in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 660–673 (1984).

D. M. Snodderly, J. D. Auran, F. C. Delori, “The macular pigment. II. Spatial distribution in primate retinas,” Invest. Ophthalmol. Visual Sci. 25, 674–685 (1984).

J. Biochem. (1)

M. Bando, A. Nakajima, K. Satoh, “Spectrophotometric estimation of 3-OH L-kynurenine O-β-glucoside in the human lens,” J. Biochem. 89, 103–109 (1981).
[PubMed]

J. Mod. Opt. (1)

J. M. Gorrand, F. C. Delori, “A model for assessment of cone directionality,” J. Mod. Opt. 44, 473–491 (1997).
[CrossRef]

J. Opt. Soc. Am. A (8)

J. Photochem. Photobiol. B (1)

J. Dillon, “The photophysics and photobiology of the eye,” J. Photochem. Photobiol. B 10, 23–40 (1991).
[CrossRef] [PubMed]

J. Physiol. (London) (2)

G. F. Cooper, J. G. Robson, “The yellow colour of the lens of man and other primates,” J. Physiol. (London) 203, 411–417 (1969).

R. A. Weale, “Age and the transmittance of the human crystalline lens,” J. Physiol. (London) 395, 577–587 (1988).

J. Vision (1)

A. Roorda, D. R. Williams, “Optical fiber properties of individual human cones,” J. Vision 2, 404–412 (2002).
[CrossRef]

Nature (London) (1)

R. van Heyningen, “Fluorescent glucoside in the human lens,” Nature (London) 230, 393–394 (1971).
[CrossRef]

Ophthalmic Res. (1)

Y. Murata, “Light absorption characteristics of the lens capsule,” Ophthalmic Res. 19, 107–112 (1987).
[CrossRef] [PubMed]

Ophthalmology (Philadelphia) (1)

T. A. Ciulla, J. Curran-Celantano, D. A. Cooper, B. R. Hammond, R. P. Danis, L. M. Pratt, K. A. Riccardi, T. G. Filloon, “Macular pigment optical density in a midwestern sample,” Ophthalmology (Philadelphia) 108, 730–737 (2001).
[CrossRef]

Opt. Lett. (1)

Optom. Vision Sci. (3)

C. A. Johnson, D. L. Howard, D. Marshall, H. Shu, “A noninvasive video-based method of measuring lens transmission properties of the human eye,” Optom. Vision Sci. 70, 944–955 (1993).
[CrossRef]

G. L. Savage, C. A. Johnson, D. L. Howard, “A comparison of noninvasive objective and subjective measurements of the optical density of human ocular media,” Optom. Vision Sci. 78, 386–395 (2001).
[CrossRef]

D. Whitaker, R. Steen, D. B. Elliott, “Light scatter in the normal young, elderly, and cataractous eye demonstrates little wavelength dependency,” Optom. Vision Sci. 70, 963–968 (1993).
[CrossRef]

Photochem. Photobiol. (1)

J. Dillon, R.-H. Wang, S. J. Atherton, “Photochemical and photophysical studies on human lens constituents,” Photochem. Photobiol. 52, 849–854 (1990).
[CrossRef] [PubMed]

Phys. Med. Biol. (2)

M. Hammer, D. Schweitzer, “Quantitative reflection spectroscopy at the human ocular fundus,” Phys. Med. Biol. 47, 179–191 (2002).
[CrossRef] [PubMed]

S. J. Preece, E. Claridge, “Monte Carlo modelling of the spectral reflectance of the human eye,” Phys. Med. Biol. 47, 2863–2877 (2002).
[CrossRef] [PubMed]

Prog. Retin. Eye Res. (1)

T. T. J. M. Berendschot, P. J. DeLint, D. van Norren, “Fundus reflectance—historical and present ideas,” Prog. Retin. Eye Res. 22, 171–200 (2003).
[CrossRef] [PubMed]

Science (1)

S. Zigman, “Eye lens color: formation and function,” Science 171, 807–809 (1971).
[CrossRef] [PubMed]

Vision Res. (15)

D. van Norren, J. J. Vos, “Spectral transmission of the human ocular media,” Vision Res. 14, 1237–1244 (1974).
[CrossRef]

S. Coren, J. S. Girgus, “Density of human lens pigmentation: in vivo measures over an extended age range,” Vision Res. 12, 343–346 (1972).
[CrossRef] [PubMed]

J. Mellerio, “Light absorption and scatter in the human lens,” Vision Res. 11, 129–141 (1971).
[CrossRef] [PubMed]

J. J. Vos, F. L. van Os, “The effect of lens density on the Stiles–Crawford effect,” Vision Res. 15, 749–751 (1975).
[CrossRef] [PubMed]

A. Stockman, L. T. Sharpe, C. Fach, “The spectral sensitivity of the human short-wavelength sensitive cones derived from thresholds and color matches,” Vision Res. 39, 2901–2927 (1999).
[CrossRef] [PubMed]

D. van Norren, L. F. Tiemeijer, “Spectral reflectance of the human eye,” Vision Res. 26, 313–320 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, D. van Norren, “Intensity and polarization of light scattered at small angles from the human fovea,” Vision Res. 26, 485–494 (1986).
[CrossRef] [PubMed]

G. J. van Blokland, “Directionality and alignment of the foveal receptors, assessed with light scattered from the human fundus in vivo,” Vision Res. 26, 495–500 (1986).
[CrossRef] [PubMed]

J.-M. Gorrand, F. Delori, “A reflectometric technique for assessing photoreceptor alignment,” Vision Res. 35, 999–1010 (1995).
[CrossRef] [PubMed]

J. van de Kraats, T. T. J. M. Berendschot, D. van Norren, “The pathways of light measured in fundus reflectometry,” Vision Res. 36, 2229–2247 (1996).
[CrossRef] [PubMed]

P. J. DeLint, T. T. J. M. Berendschot, D. van Norren, “Local photoreceptor alignment measured with a scanning laser ophthalmoscope,” Vision Res. 37, 243–248 (1997).
[CrossRef] [PubMed]

T. J. T. P. van den Berg, J. K. IJspeert, “Light scattering in donor lenses,” Vision Res. 35, 169–177 (1995).
[CrossRef] [PubMed]

J. S. Werner, S. K. Donnelly, R. Kliegl, “Aging and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering,” Vision Res. 27, 257–268 (1987).
[CrossRef] [PubMed]

B. R. Wooten, G. A. Geri, “Psychophysical determination of intraocular light scatter as a function of wavelength,” Vision Res. 27, 1291–1298 (1987).
[CrossRef] [PubMed]

T. J. T. P. van den Berg, K. E. W. P. Tan, “Light transmittance of the human cornea from 320 to 700 nm for different ages,” Vision Res. 34, 1453–1456 (1994).
[CrossRef] [PubMed]

Other (10)

J. Marshall, “Ageing changes in human cones,” in International Congress Series, K. Shimizu, J. A. Oosterhuis, eds. (Excerpta Media, Amsterdam, 1979), pp. 375–378.

Alcon Nederland BV, Product Information Booklet (Alcon Nederland BV, Avelingen-West 5, 4202 MS, Gorinchem, The Netherlands, 2002).

E. A. Boettner, Spectral Transmission of the Eye [Report of the University of Michigan, contract AF41(609)-2966, Ann Arbor, Mich., 1967].

G. Wyszecki, W. S. Stiles, Color Science: Concepts and Methods, Quantitative Data and Formulae, 2nd ed. (Wiley, New York, 1982).

W. H. Press, B. P. Flannery, S. A. Teukolsky, W. T. Vetterling, Numerical Recipes in Pascal (Cambridge U. Press, Cambridge, UK, 1989).

P. L. Walraven, , Draft 7 (personal communication, 2003) (plwalraven@hotmail.com).

J. J. Vos, Literature Review of Human Macular Absorption in the Visible and Its Consequences for the Cone Receptor Primaries (National Organization for Applied Scientific Research in The Netherlands, Institute for Perception RVO-TNO, Soesterberg, The Netherlands, 1972).

J. Krauskopf, “Some experiments with a photoelectric ophthalmoscope,” in Performance of the Eye at Low Luminances, M. A. Bouman, J. J. Vos, eds. (Excerpta Medica, Amsterdam, 1966), pp. 171–181.

K. E. W. P. Tan, “Vision in the ultraviolet,” Ph.D thesis (Universiteit Utrecht, Utrecht, The Netherlands, 1971).

S. A. Burns, J. C. He, F. C. Delori, “Do the cones see light scattered from the deeper retinal layers?” in Vision Science and Its Applications, Vol. 1 of 1997 OSA Technical Digest Series (Optical Society of America, Washington D.C., 1997), pp. 94–97.

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Figures (8)

Fig. 1
Fig. 1

(A) Optical density spectra of macular pigment, taken from the literature and normalized to unity at 460 nm. Filled triangles, literature review by Vos23 (1972); squares, literature review by Wyszecki and Stiles20 (1982); open triangles, data by Stockman et al.24 (1999); circles, data by Brown, cited by Delori et al.25 (2001). The solid curve was calculated with the template [Eq. (3)] by Walraven.22 Note that the spectra are not independent in that the reviews by Vos and by Wyszecki and Stiles largely rely on the same data. (B) The optical density spectrum of an intraocular lens adapted from an Alcon product information booklet.26 The solid curve produced with Eq. (4) closely follows the discrete data points.

Fig. 2
Fig. 2

(A–E) Top: Amplitude of the directional reflectance converted into a density Ddir(λ) for the pseudophakic subjects. Symbols represent the mean density; error bars indicate the total errors in the mean. The subjects’ ages and the number of averaged spectra are indicated. Solid curves depict model fits of Eq. (5) to the data. Bottom: The differences with the fit at an enhanced scale. (F) The same results for the mean of the five subjects’ data. All the six spectra reflect the optical density spectrum of macular pigment; cf. Fig. 1A.

Fig. 3
Fig. 3

(A) Mean optical density Ddir(λ) for ten young and ten aged subjects. The error bars, not discernable in most cases, indicate the total errors in the mean. The data were not shifted vertically; the optical density in aged eyes is larger at longer wavelengths as well. The rise in optical density from 450 to 420 nm is due to absorption in the crystalline lens. The dashed curves depict the best fit with the Pokorny et al.31 lens aging model. Solid curves represent the simultaneous fit of Eq. (8) to both data sets in order to derive two new templates for the absorption in the lens. (B, C) The differences between the mean aged and young data and the model fit based on the Pokorny et al.31 lens aging model (triangles) and the model based on Eq. (8) (squares). The new templates improved the fit to the data.

Fig. 4
Fig. 4

(A) Optical density of 3-HKG, normalized to unity at 370 nm; crosses, Dillon40 (1984); squares, Dillon et al.36 (1990); circles, Heckathorn et al.41 (2001). The diamonds represent the spectrum of 3-hydroxykynurenine (3-HK) by Stutchbury and Truscott,44 (1993), which is similar to the 3-HKG spectra. The Dillon et al. (1990) spectrum36 was first reported representing 3-HK, but in 1991 Dillon presented the same data as a 3-HKG spectrum.43 The spectrum of the eye of a young monkey (open triangles) by Gaillard et al.38 (2000) is similar to the 3-HK and 3-HKG spectra. The solid curve depicts a single Gaussian fitted to the Heckathorn et al. 3-HKG data.41 The filled triangles represent the Pokorny et al. nonaging template.31 It has the same shape as the 3-HKG spectra. (B) Various suggested optical density spectra for the pigments accumulating with age, normalized to unity at 420 nm; squares, Dillon et al.36 (1990); crosses, D́illon et al.45 (1999); triangles, Gaillard et al.38 (2000). The circles depict the Pokorny et al.31 template affected by aging. The curves drawn through the above data sets represent a fit with the tail of a single Gaussian or the sum of two Gaussians centered at 320 nm. The thick solid curve depicts the newly proposed aged template [Eq. (9)].

Fig. 5
Fig. 5

(A–F) Top: Amplitude of the directional fundus reflectance, converted into a density, versus wavelength for a sample of six normal subjects spanning the full age range. The symbols represent the mean density within a series of five measurements. Error bars indicate the total errors in the mean. Ages are indicated above each spectrum. The solid curves depict model fits of Eq. (10) to the data. Compared with Fig. 2, the spectra steeply rise below 450 nm. The additional optical density is caused by absorption in the crystalline lens. Bottom: The differences between the data and the model fit.

Fig. 6
Fig. 6

Densities of (A) the young and (B) the aged components and (C) the total lens density versus age at 420 nm. The error bars indicate the statistical error estimated in the model fits. The solid curves depict linear regression lines; their parameters are given in Table 2. The density of the young component decreased with age. The density of the aged component strongly increased with age. The total optical density of the lens slightly increased with age.

Fig. 7
Fig. 7

(A) Macular pigment density and (B) the reflectivity of the receptors versus age. The circles represent individual estimates for the 39 normal subjects; the crosses depict data obtained in the pseudophakes. The error bars are smaller than the symbols in most cases. The solid curves depict a linear regression to the normal subjects’ data; parameters are given in Table 2. The receptor reflectance strongly decreased with age.

Fig. 8
Fig. 8

Comparison of lens optical density spectra for (A) observers aged approximately 20 years and (B) observers aged approximately 50 years. Solid curves represent the present model calculated with Eq. (11). Data obtained with the Purkinje image method: open triangles, Johnson et al.50 (1993), age groups 21–30 and 41–50 years; filled triangles, Savage et al.51 (2001), for observers with average ages of 24 and 50 years. Psychophysical data: circles, the Pokorny31 et al. (1987) model; dashed curves, the Savage33 et al. (1993) model; solid squares, Savage et al.51 (2001), in the same observers with average ages of 24 and 50 years that provided the objective data represented by the solid triangles.

Tables (2)

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Table 1 Model Fit Parameters and Their Statistical Errors Estimated from the Fit

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Table 2 Parameters Derived from Linear Regression with Age

Equations (11)

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R(x)=B+Adir10-ρ(x-xC)2,
Adir(λ)=Rrecep(λ)10-2[Dmacαmac(λ)+Dmediaαmedia(λ)],
αmac(λ)=fnorm{0.32 exp[-0.0012(436-λ)2]+0.32 exp[-0.0012(480-λ)2]-0.123 exp[-0.0012(458-λ)2]+0.12042 exp[-0.006(457-λ)2]},
DIOLαIOL(λ)=0.0838 exp{-[(λ-420)/36.6]}+8.55 exp{-[(λ-380)/12.9]2}.
Ddir(λ)=Cr+2 Dmacαmac(λ)+2 DIOLαIOL(λ),
Ddir(λ)=Cr+2 Dmacαmac(λ)+2 Dlens-aαlens-a(λ)+2 0.30αlens-na(λ),
αyoung(λ)=6.09 exp{-[(λ-370)/37.2]2}.
Ddir(λ)=Cr+2 Dmacαmac(λ)+2 Dyoungαyoung(λ)+2 Dagedexp{[(320-420)/Δ]2}×exp{-[(λ-320)/Δ]2},
αaged(λ)=5.65 exp{-[(λ-320)/76.0]2}.
Ddir(λ)=Cr+2 Dmacαmac(λ)+2 Dyoungαyoung(λ)+2 Dagedαaged(λ),
Dmedia(λ, A)=(0.42-0.0064 A)αyoung(λ)+(-0.08+0.0094 A)αaged(λ),

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