Abstract

Cone oil droplets are highly retractile, optically dense spheres located in front of the cone outer segment. Because of their refractile strength, absorbance, and size, these spherical oil droplets have a pronounced influence on the light reaching the photopigment. We used electromagnetic scattering theory to interrelate these factors, and we calculated the light energy distribution around cone oil droplets modeled as isolated spheres. The necessary electromagnetic parameters were determined from anatomical and optical measurements in the turtle retina. Cone and oil-droplet dimensions were determined with a light microscope. The index-of-refraction values for the red, yellow, orange, and clear oil droplets were measured to be 1.69, 1.55, 1.51, and 1.48, respectively, by the technique of immersion matching. The application of the electromagnetic scattering theory suggests that oil droplets significantly increase cone sensitivity. Cone action spectra predicted by this analysis show that the largest increases o ccur around the absorption maximum of each cone's photopigment.

© 1983 Optical Society of America

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  1. G. L. Walls, The Vertebrate Eye and Its Adaptive Radiation (Cranbrook Institute of Science, Bloomfield Hills, Mich, 1942).
  2. P. A. Liebman and A. M. Granda, "Super dense carotenoid spectra resolved in single cone oil droplets," Nature 253,370–372 (1975).
  3. H. F. Leeper, "Horizontal cells of the turtle retina: analysis of interconnections between photoreceptor cells and horizontal cells by light microscopy," J. Comp. Neurol. 182, 795–810 (1978).
  4. D. A. Baylor and R. Fettiplace, "Light path and photon capture in turtle photoreceptors," J. Physiol. 248, 433–464 (1975).
  5. M. L. Wolbarsht, "The function of intraocular color filters," Fed. Proc. 35, 44–50 (1976).
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  7. J. K. Bowmaker and A. Knowles, "The visual pigments and oil droplets of the chicken retina," Vision Res. 17, 755–764 (1977).
  8. G. H. Jacobs, Comparative Color Vision (Academic, New York, 1981).
  9. J. A. Stratton, Electromagnetic Theory (McGraw-Hill, New York, 1941).
  10. H. C. Van de Hulst, Light Scattering by Small Particles (Wiley, New York, 1957).
  11. R. F. Harrington, Time-Harmonic Electromagnetic Fields (McGraw-Hill, New York, 1961).
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  30. M. A. Ali, "Les reponses retinometrices: caracteres et mecanismes," Vision Res. 11, 1225–1288 (1971).

1982 (1)

H. Kolb and J. Jones, "Light and electron microscopy of the photoreceptors in the retina of the red eared slider, Pseudemys scripta elegans," J. Comp. Neurol. 209, 331–338 (1982).

1978 (1)

H. F. Leeper, "Horizontal cells of the turtle retina: analysis of interconnections between photoreceptor cells and horizontal cells by light microscopy," J. Comp. Neurol. 182, 795–810 (1978).

1977 (1)

J. K. Bowmaker and A. Knowles, "The visual pigments and oil droplets of the chicken retina," Vision Res. 17, 755–764 (1977).

1976 (2)

M. L. Wolbarsht, "The function of intraocular color filters," Fed. Proc. 35, 44–50 (1976).

W. S. Jaggers and P. A. Liebman, "Anomalous dispersion of rhodopsin in rod outer segments of the frog," J. Opt. Soc. Am. 66, 56–59 (1976).

1975 (2)

P. A. Liebman and A. M. Granda, "Super dense carotenoid spectra resolved in single cone oil droplets," Nature 253,370–372 (1975).

D. A. Baylor and R. Fettiplace, "Light path and photon capture in turtle photoreceptors," J. Physiol. 248, 433–464 (1975).

1973 (2)

D. A. Baylor and A. L. Hodgkin, "Detection and resolution of visual stimuli by turtle photoreceptors," J. Physiol. 234, 163–198 (1973).

A. M. Snyder and C. Pask, "The Stiles—Crawford effect—explanation and consequences," Vision Res. 13, 1115–1137 (1973).

1971 (2)

M. A. Ali, "Les reponses retinometrices: caracteres et mecanismes," Vision Res. 11, 1225–1288 (1971).

P. A. Liebman and A. M. Granda, "Microspectrophotometric measurements of visual pigments in two species of turtle, Pseudemys scripta and Chelonia mydas," Vision Res. 11, 105–114 (1971).

1970 (1)

A. M. Granda and K. W. Haden, "Retinal oil globule counts and distributions in two species of turtle: Pseudemys scripta elegans (Wied) and Chelonia mydas mydas (Linnaeus)," Vision Res. 10, 79–84 (1970).

1969 (1)

K. T. Brown, "A linear area centralis extending across the turtle retina and stabilized to the horizon by non-visual cues," Vision Res. 9, 1053–1062 (1969).

1967 (1)

1963 (2)

G. K. Strother, "Absorption spectra of retinal oil globules in turkey, turtle and pigeon," Expl. Cell Res. 29, 349–355 (1963).

J. M. Enoch, "Optical role of retinal receptors," J. Opt. Soc. Am. 53, 71–85 (1963).

1961 (1)

1960 (1)

1957 (1)

R. L. Sidman, "The structure and concentration of solids in photoreceptor cells studied by refractometry and interference microscopy," J. Biophys. Biochem. Cytol. 3, 15–33 (1957).

1956 (1)

K. Tansley and B. K. Johnson, "The cones of the grass snake's eyes," Nature 178, 1285–1286 (1956).

Ali, M. A.

M. A. Ali, "Les reponses retinometrices: caracteres et mecanismes," Vision Res. 11, 1225–1288 (1971).

Baylor, D. A.

D. A. Baylor and R. Fettiplace, "Light path and photon capture in turtle photoreceptors," J. Physiol. 248, 433–464 (1975).

D. A. Baylor and A. L. Hodgkin, "Detection and resolution of visual stimuli by turtle photoreceptors," J. Physiol. 234, 163–198 (1973).

Bennett, A. H.

A. H. Bennett, H. Jupnik, H. Osterberg, and O. W. Richards, Phase Microscopy: Principles and Applications (Wiley, New York, 1951).

Bowmaker, J. K.

J. K. Bowmaker and A. Knowles, "The visual pigments and oil droplets of the chicken retina," Vision Res. 17, 755–764 (1977).

Brown, K. T.

K. T. Brown, "A linear area centralis extending across the turtle retina and stabilized to the horizon by non-visual cues," Vision Res. 9, 1053–1062 (1969).

Detweiler, S. R.

S. R. Detweiler, Vertebrate Photoreceptors (Macmillan, New York, 1943).

Ditchburn, R. W.

R. W. Ditchburn, Light (Interscience, New York, 1953).

Enoch, J. M.

Fettiplace, R.

D. A. Baylor and R. Fettiplace, "Light path and photon capture in turtle photoreceptors," J. Physiol. 248, 433–464 (1975).

Granda, A. M.

P. A. Liebman and A. M. Granda, "Super dense carotenoid spectra resolved in single cone oil droplets," Nature 253,370–372 (1975).

P. A. Liebman and A. M. Granda, "Microspectrophotometric measurements of visual pigments in two species of turtle, Pseudemys scripta and Chelonia mydas," Vision Res. 11, 105–114 (1971).

A. M. Granda and K. W. Haden, "Retinal oil globule counts and distributions in two species of turtle: Pseudemys scripta elegans (Wied) and Chelonia mydas mydas (Linnaeus)," Vision Res. 10, 79–84 (1970).

Haden, K. W.

A. M. Granda and K. W. Haden, "Retinal oil globule counts and distributions in two species of turtle: Pseudemys scripta elegans (Wied) and Chelonia mydas mydas (Linnaeus)," Vision Res. 10, 79–84 (1970).

Harrington, R. F.

R. F. Harrington, Time-Harmonic Electromagnetic Fields (McGraw-Hill, New York, 1961).

Hodgkin, A. L.

D. A. Baylor and A. L. Hodgkin, "Detection and resolution of visual stimuli by turtle photoreceptors," J. Physiol. 234, 163–198 (1973).

Jacobs, G. H.

G. H. Jacobs, Comparative Color Vision (Academic, New York, 1981).

Jaggers, W. S.

Johnson, B. K.

K. Tansley and B. K. Johnson, "The cones of the grass snake's eyes," Nature 178, 1285–1286 (1956).

Jones, J.

H. Kolb and J. Jones, "Light and electron microscopy of the photoreceptors in the retina of the red eared slider, Pseudemys scripta elegans," J. Comp. Neurol. 209, 331–338 (1982).

Jupnik, H.

A. H. Bennett, H. Jupnik, H. Osterberg, and O. W. Richards, Phase Microscopy: Principles and Applications (Wiley, New York, 1951).

Kerker, M.

M. Kerker, The Scattering of Light and Other Electromagnetic Radiation (Academic, New York, 1969).

Knowles, A.

J. K. Bowmaker and A. Knowles, "The visual pigments and oil droplets of the chicken retina," Vision Res. 17, 755–764 (1977).

Kolb, H.

H. Kolb and J. Jones, "Light and electron microscopy of the photoreceptors in the retina of the red eared slider, Pseudemys scripta elegans," J. Comp. Neurol. 209, 331–338 (1982).

Leeper, H. F.

H. F. Leeper, "Horizontal cells of the turtle retina: analysis of interconnections between photoreceptor cells and horizontal cells by light microscopy," J. Comp. Neurol. 182, 795–810 (1978).

Liebman, P. A.

W. S. Jaggers and P. A. Liebman, "Anomalous dispersion of rhodopsin in rod outer segments of the frog," J. Opt. Soc. Am. 66, 56–59 (1976).

P. A. Liebman and A. M. Granda, "Super dense carotenoid spectra resolved in single cone oil droplets," Nature 253,370–372 (1975).

P. A. Liebman and A. M. Granda, "Microspectrophotometric measurements of visual pigments in two species of turtle, Pseudemys scripta and Chelonia mydas," Vision Res. 11, 105–114 (1971).

P. A. Liebman, Handbook of Sensory Physiology VII/1 (Springer-Verlag, Berlin, 1972).

Osterberg, H.

A. H. Bennett, H. Jupnik, H. Osterberg, and O. W. Richards, Phase Microscopy: Principles and Applications (Wiley, New York, 1951).

Pask, C.

A. M. Snyder and C. Pask, "The Stiles—Crawford effect—explanation and consequences," Vision Res. 13, 1115–1137 (1973).

Richards, O. W.

A. H. Bennett, H. Jupnik, H. Osterberg, and O. W. Richards, Phase Microscopy: Principles and Applications (Wiley, New York, 1951).

Sidman, R. L.

R. L. Sidman, "The structure and concentration of solids in photoreceptor cells studied by refractometry and interference microscopy," J. Biophys. Biochem. Cytol. 3, 15–33 (1957).

Snyder, A. M.

A. M. Snyder and C. Pask, "The Stiles—Crawford effect—explanation and consequences," Vision Res. 13, 1115–1137 (1973).

Stratton, J. A.

J. A. Stratton, Electromagnetic Theory (McGraw-Hill, New York, 1941).

Strother, G. K.

G. K. Strother, "Absorption spectra of retinal oil globules in turkey, turtle and pigeon," Expl. Cell Res. 29, 349–355 (1963).

Tansley, K.

K. Tansley and B. K. Johnson, "The cones of the grass snake's eyes," Nature 178, 1285–1286 (1956).

Van de Hulst, H. C.

H. C. Van de Hulst, Light Scattering by Small Particles (Wiley, New York, 1957).

Walls, G. L.

G. L. Walls, The Vertebrate Eye and Its Adaptive Radiation (Cranbrook Institute of Science, Bloomfield Hills, Mich, 1942).

Wolbarsht, M. L.

M. L. Wolbarsht, "The function of intraocular color filters," Fed. Proc. 35, 44–50 (1976).

Expl. Cell Res. (1)

G. K. Strother, "Absorption spectra of retinal oil globules in turkey, turtle and pigeon," Expl. Cell Res. 29, 349–355 (1963).

Fed. Proc. (1)

M. L. Wolbarsht, "The function of intraocular color filters," Fed. Proc. 35, 44–50 (1976).

J. Biophys. Biochem. Cytol. (1)

R. L. Sidman, "The structure and concentration of solids in photoreceptor cells studied by refractometry and interference microscopy," J. Biophys. Biochem. Cytol. 3, 15–33 (1957).

J. Comp. Neurol. (1)

H. F. Leeper, "Horizontal cells of the turtle retina: analysis of interconnections between photoreceptor cells and horizontal cells by light microscopy," J. Comp. Neurol. 182, 795–810 (1978).

J. Opt. Soc. Am. (5)

J. Physiol. (2)

D. A. Baylor and R. Fettiplace, "Light path and photon capture in turtle photoreceptors," J. Physiol. 248, 433–464 (1975).

D. A. Baylor and A. L. Hodgkin, "Detection and resolution of visual stimuli by turtle photoreceptors," J. Physiol. 234, 163–198 (1973).

Nature (2)

P. A. Liebman and A. M. Granda, "Super dense carotenoid spectra resolved in single cone oil droplets," Nature 253,370–372 (1975).

K. Tansley and B. K. Johnson, "The cones of the grass snake's eyes," Nature 178, 1285–1286 (1956).

Vision Res. (4)

M. A. Ali, "Les reponses retinometrices: caracteres et mecanismes," Vision Res. 11, 1225–1288 (1971).

A. M. Snyder and C. Pask, "The Stiles—Crawford effect—explanation and consequences," Vision Res. 13, 1115–1137 (1973).

K. T. Brown, "A linear area centralis extending across the turtle retina and stabilized to the horizon by non-visual cues," Vision Res. 9, 1053–1062 (1969).

J. K. Bowmaker and A. Knowles, "The visual pigments and oil droplets of the chicken retina," Vision Res. 17, 755–764 (1977).

Other (13)

G. H. Jacobs, Comparative Color Vision (Academic, New York, 1981).

J. A. Stratton, Electromagnetic Theory (McGraw-Hill, New York, 1941).

H. C. Van de Hulst, Light Scattering by Small Particles (Wiley, New York, 1957).

R. F. Harrington, Time-Harmonic Electromagnetic Fields (McGraw-Hill, New York, 1961).

M. Kerker, The Scattering of Light and Other Electromagnetic Radiation (Academic, New York, 1969).

G. L. Walls, The Vertebrate Eye and Its Adaptive Radiation (Cranbrook Institute of Science, Bloomfield Hills, Mich, 1942).

P. A. Liebman and A. M. Granda, "Microspectrophotometric measurements of visual pigments in two species of turtle, Pseudemys scripta and Chelonia mydas," Vision Res. 11, 105–114 (1971).

A. M. Granda and K. W. Haden, "Retinal oil globule counts and distributions in two species of turtle: Pseudemys scripta elegans (Wied) and Chelonia mydas mydas (Linnaeus)," Vision Res. 10, 79–84 (1970).

H. Kolb and J. Jones, "Light and electron microscopy of the photoreceptors in the retina of the red eared slider, Pseudemys scripta elegans," J. Comp. Neurol. 209, 331–338 (1982).

P. A. Liebman, Handbook of Sensory Physiology VII/1 (Springer-Verlag, Berlin, 1972).

R. W. Ditchburn, Light (Interscience, New York, 1953).

S. R. Detweiler, Vertebrate Photoreceptors (Macmillan, New York, 1943).

A. H. Bennett, H. Jupnik, H. Osterberg, and O. W. Richards, Phase Microscopy: Principles and Applications (Wiley, New York, 1951).

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