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Journal of the Optical Society of America A

Journal of the Optical Society of America A


  • Editor: Franco Gori
  • Vol. 29, Iss. 2 — Feb. 1, 2012
  • pp: A19–A26

Mesopic rod and S-cone interactions revealed by modulation thresholds

Andrew J. Zele, Jan Kremers, and Beatrix Feigl  »View Author Affiliations

JOSA A, Vol. 29, Issue 2, pp. A19-A26 (2012)

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We analyzed mesopic rod and S-cone interactions in terms of their contributions to the blue–yellow opponent pathway. Stimuli were generated using a four-primary colorimeter. Mixed rod and S-cone modulation thresholds (constant L-, M-cone excitation) were measured as a function of their phase difference. Modulation amplitude was equated using threshold units and contrast ratios. This study identified three interaction types: (1) a linear and antagonistic rod:S-cone interaction, (2) probability summation, and (3) a previously unidentified mutual nonlinear reinforcement. Linear rod:S-cone interactions occur within the blue–yellow opponent pathway. Probability summation involves signaling by different postreceptoral pathways. The origin of the nonlinear reinforcement is possibly at the photoreceptors.

© 2012 Optical Society of America

OCIS Codes
(330.0330) Vision, color, and visual optics : Vision, color, and visual optics
(330.1720) Vision, color, and visual optics : Color vision
(330.5510) Vision, color, and visual optics : Psychophysics

ToC Category:
Mesopic and peripheral color vision

Original Manuscript: August 4, 2011
Revised Manuscript: October 25, 2011
Manuscript Accepted: November 1, 2011
Published: December 22, 2011

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Vol. 7, Iss. 4 Virtual Journal for Biomedical Optics

Andrew J. Zele, Jan Kremers, and Beatrix Feigl, "Mesopic rod and S-cone interactions revealed by modulation thresholds," J. Opt. Soc. Am. A 29, A19-A26 (2012)

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  1. N. W. Daw, E. J. Jensen, and W. J. Bunken, “Rod pathways in the mammalian retinae,” Trends Neurosci. 13, 110–115 (1990). [CrossRef] [PubMed]
  2. G. D. Field, M. Greschner, J. L. Gauthier, C. Rangel, J. Shlens, A. Sher, D. W. Marshak, A. M. Litke, and E. J. Chichilnisky, “High-sensitivity rod photoreceptor input to the blue-yellow color opponent pathway in macaque retina,” Nat. Neurosci. 12, 1159–1164 (2009). [CrossRef] [PubMed]
  3. J. D. Crook, C. M. Davenport, B. B. Peterson, O. S. Packer, P. B. Detwiler, and D. M. Dacey, “Parallel ON and OFF cone bipolar inputs establish spatially coextensive receptive field structure of blue-yellow ganglion cells in primate retina,” J. Neurosci. 29, 8372–8387 (2009). [CrossRef] [PubMed]
  4. V. Virsu and B. B. Lee, “Light adaptation in cells of macaque lateral geniculate nucleus and its relation to human light adaptation,” J. Neurophysiol. 50, 864–878 (1983). [PubMed]
  5. B. B. Lee, V. C. Smith, J. Pokorny, and J. Kremers, “Rod inputs to macaque ganglion cells,” Vis. Res. 37, 2813–2828 (1997). [CrossRef]
  6. T. Wiesel and D. H. Hubel, “Spatial and chromatic interactions in the lateral geniculate body of the rhesus monkey,” J. Neurophysiol. 29, 1115–1156 (1966). [PubMed]
  7. D. Cao, B. B. Lee, and H. Sun, “Combination of rod and cone inputs in parasol ganglion cells of the magnocellular pathway,” J. Vis. 10, 4 (2010). [CrossRef] [PubMed]
  8. V. Virsu, B. B. Lee, and O. D. Creutzfeldt, “Dark adaptation and receptive field organisation of cells in the cat lateral geniculate nucleus,” Exp. Brain Res. 27, 35–50 (1977). [CrossRef] [PubMed]
  9. B. B. Lee, J. Pokorny, V. C. Smith, P. R. Martin, and A. Valberg, “Luminance and chromatic modulation sensitivity of macaque ganglion cells and human observers,” J. Opt. Soc. Am. A 7, 2223–2236 (1990). [CrossRef] [PubMed]
  10. K. Purpura, D. Tranchina, E. Kaplan, and R. M. Shapley, “Light adaptation in the primate retina: analysis of changes in gain and dynamics of monkey retinal ganglion cells,” Vis. Neurosci. 4, 75–93 (1990). [CrossRef] [PubMed]
  11. S. Weiss, J. Kremers, and J. Maurer, “Interaction between rod and cone signals in responses of lateral geniculate neurons in dichromatic marmosets (Callithrix jacchus),” Vis. Neurosci. 15, 931–943 (1998). [CrossRef] [PubMed]
  12. D. M. Dacey, “Morphology of a small-field bistratified ganglion cell type in the macaque and human retina,” Vis. Neurosci. 10, 1081–1098 (1993). [CrossRef] [PubMed]
  13. D. M. Dacey and B. B. Lee, “The ‘blue-on’ opponent pathway in primate retina originates from a distinct bistratified ganglion cell type,” Nature 367, 731–735 (1994). [CrossRef] [PubMed]
  14. G. D. Field, J. L. Gauthier, A. Sher, M. Greschner, T. A. Machado, L. H. Jepson, J. Shlens, D. E. Gunning, K. Mathieson, W. Dabrowski, L. Paninski, A. M. Litke, and E. J. Chichilnisky, “Functional connectivity in the retina at the resolution of photoreceptors,” Nature 467, 673–677 (2010). [CrossRef] [PubMed]
  15. B. B. Lee, P. R. Martin, and U. Grunert, “Retinal connectivity and primate vision,” Prog. Retin. Eye Res. 29, 622–639 (2010). [CrossRef] [PubMed]
  16. T. Yeh, B. B. Lee, and J. Kremers, “The temporal response of ganglion cells of the macaque retina to cone-specific modulation,” J. Opt. Soc. Am. A 12, 456–464 (1995). [CrossRef]
  17. O. S. Packer, J. Verweij, P. H. Li, J. L. Schnapf, and D. M. Dacey, “Blue-yellow opponency in primate S cone photoreceptors,” J. Neurosci. 30, 568–572 (2010). [CrossRef] [PubMed]
  18. D. Cao, J. Pokorny, and V. C. Smith, “Matching rod percepts with cone stimuli,” Vis. Res. 45, 2119–2128 (2005). [CrossRef] [PubMed]
  19. D. Cao, J. Pokorny, V. C. Smith, and A. J. Zele, “Rod contributions to color perception: linear with rod contrast,” Vision Res. 48, 2586–2592 (2008). [CrossRef] [PubMed]
  20. D. Cao, A. J. Zele, and J. Pokorny, “Chromatic discrimination in the presence of incremental and decremental rod pedestals,” Vis. Neurosci. 25, 399–404 (2008). [CrossRef] [PubMed]
  21. S. L. Buck, E. Sanocki, and R. Knight, “Do rod signals add with S cone signals in increment detection?” in Colour Vision Deficiencies, C.R.Cavonius, ed. (Kluwer, 1997), pp. 451–458. [CrossRef]
  22. H. Sun, J. Pokorny, and V. C. Smith, “Brightness induction from rods,” J. Vis. 1, 32–41 (2001). [CrossRef]
  23. R. Knight, S. L. Buck, G. A. Fowler, and A. Nguyen, “Rods affect S-cone discrimination on the Farnsworth–Munsell 100-hue test,” Vis. Res. 38, 3477–3481 (1998). [CrossRef]
  24. J. Pokorny, M. Lutze, D. Cao, and A. J. Zele, “The color of night: Surface color perception under dim illuminations,” Vis. Neurosci. 23, 525–530 (2006). [CrossRef] [PubMed]
  25. E. N. Willmer, “Low threshold rods and the perception of blue,” J. Physiol. 111, 10–20, Abstract 17P (1949).
  26. I. Lie, “Dark adaptation and the photochromatic interval,” Doc. Ophthalmol. 17, 411–510 (1963). [CrossRef] [PubMed]
  27. P. W. Trezona, “Rod participation in the ‘blue’ mechanism and its effect on colour matching,” Vis. Res. 10, 317–332 (1970). [CrossRef] [PubMed]
  28. U. Stabell and B. Stabell, “Chromatic rod vision. II. Wavelength of pre-stimulation varied,” Scand. J. Psychol. 12, 282–288(1971). [CrossRef] [PubMed]
  29. S. L. Buck, R. Knight, G. Fowler, and B. Hunt, “Rod influence on hue-scaling functions,” Vis. Res. 38, 3259–3263 (1998). [CrossRef]
  30. G. Lange, N. Denny, and T. E. Frumkes, “Suppressive rod–cone interactions: evidence for separate retinal (temporal) and extraretinal (spatial) mechanisms in achromatic vision,” J. Opt. Soc. Am. A 14, 2487–2498 (1997). [CrossRef]
  31. D. Cao, A. J. Zele, and J. Pokorny, “Dark-adapted rod suppression of cone flicker detection: evaluation of receptoral and postreceptoral interactions,” Vis. Neurosci. 23, 531–537 (2006). [CrossRef] [PubMed]
  32. J. Kremers and S. Meierkord, “Rod–cone-interactions in deuteranopic observers: models and dynamics,” Vis. Res. 39, 3372–3385 (1999). [CrossRef]
  33. H. Sun, J. Pokorny, and V. C. Smith, “Rod-cone interactions assessed in inferred magnocellular and parvocellular postreceptoral pathways,” J. Vis. 1, 42–54 (2001). [CrossRef]
  34. A. J. Zele, D. Cao, and J. Pokorny, “Rod-cone interactions and the temporal impulse response of the cone pathway,” Vis. Res. 48, 2593–2598 (2008). [CrossRef] [PubMed]
  35. A. J. Zele and A. J. Vingrys, “Defining the detection mechanisms for symmetric and rectified flicker stimuli,” Vis. Res. 47, 2700–2713 (2007). [CrossRef] [PubMed]
  36. W. Nagel, “Appendix: adaptation, twilight vision and the duplicity theory,” in Helmholtz’s Treatise on Physiological Optics. Translated from the Third German Edition by J. P. C. Southall, 3rd German ed. (Optical Society of America, 1924), pp. 313–343.
  37. S. L. Buck, “What is the hue of rod vision?” Color Res. Appl. 26, S57–S59 (2001). [CrossRef]
  38. J. Purkinje, Beobachtungen und Versuche zur Physiologie der Sinne. Neue Beiträge zur Kenntniss des Sehens in subjectiver Hinsicht (Reimer, 1825).
  39. E. A. Hough, “The spectral sensitivity functions for parafoveal vision,” Vis. Res. 8, 1423–1430 (1968). [CrossRef] [PubMed]
  40. E. A. Hough and K. H. Ruddock, “The parafoveal visual response of a tritanope and an interpretation of the Vλ sensitivity functions of mesopic vision,” Vis. Res. 9, 935–946 (1969). [CrossRef] [PubMed]
  41. E. A. Hough and K. H. Ruddock, “The Purkinje shift,” Vis. Res. 9, 313–315 (1969). [CrossRef] [PubMed]
  42. Commission Internationale de l’Éclairage, “Mesopic photometry: history, special problems and practical solutions. Publ. CIE No. 81,” TC 1-01 (Bureau Central de la Commission Internationale de l’Éclairage, 1989).
  43. H. R. Blackwell and O. M. Blackwell, “Rod and cone receptor mechanisms in typical and atypical congenital achromatopsia,” Vis. Res. 1, 62–107 (1961). [CrossRef]
  44. J. Pokorny, V. C. Smith, and R. Swartley, “Threshold measurements of spectral sensitivity in a blue monocone monochromat,” Invest. Ophthalmol. Visual Sci. 9, 807–813 (1970).
  45. M. Alpern, G. B. Lee, and B. E. Spivey, “Pi-1 cone monochromatism,” Arch. Ophthalmol. 74, 334–337 (1965). [CrossRef] [PubMed]
  46. F. Naarendorp, K. S. Rice, and P. A. Sieving, “Summation of rod and S cone signals at threshold in human observers,” Vis. Res. 36, 2681–2688 (1996). [CrossRef] [PubMed]
  47. J. L. Schnapf, B. J. Nunn, M. Meister, and D. A. Baylor, “Visual transduction in cones of the monkey Macaca fascicularis,” J. Physiol. 427, 681–713 (1990). [PubMed]
  48. V. C. Smith, J. Pokorny, B. B. Lee, and D. M. Dacey, “Sequential processing in vision: the interaction of sensitivity regulation and temporal dynamics,” Vis. Res. 48, 2649–2656 (2008). [CrossRef] [PubMed]
  49. E. Kaplan, “The M, P, and K pathways of the primate visual system,” in The Visual Neurosciences, L.M.Chalupa and J.S.Werner, eds. (MIT, 2004), pp. 481–493.
  50. R. M. Boynton, M. Ikeda, and W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vis. Res. 4, 87–117 (1964). [CrossRef] [PubMed]
  51. T. J. T. P. van den Berg and H. Spekreijse, “Interaction between rod and cone signals studied with temporal sine wave stimulation,” J. Opt. Soc. Am. 67, 1210–1217 (1977). [CrossRef] [PubMed]
  52. J. Pokorny, H. Smithson, and J. Quinlan, “Photostimulator allowing independent control of rods and the three cone types,” Vis. Neurosci. 21, 263–267 (2004). [CrossRef] [PubMed]
  53. A. G. Shapiro, J. Pokorny, and V. C. Smith, “Cone-rod receptor spaces, with illustrations that use CRT phosphor and light-emitting-diode spectra,” J. Opt. Soc. Am. A 13, 2319–2328(1996). [CrossRef]
  54. M. J. Puts, J. Pokorny, J. Quinlan, and L. Glennie, “Audiophile hardware in vision science; the soundcard as a digital to analog converter,” J. Neurosci. Methods 142, 77–81 (2005). [CrossRef] [PubMed]
  55. W. H. Swanson, T. Ueno, V. C. Smith, and J. Pokorny, “Temporal modulation sensitivity and pulse detection thresholds for chromatic and luminance perturbations,” J. Opt. Soc. Am. A 4, 1992–2005 (1987). [CrossRef] [PubMed]
  56. A. J. Zele, D. Cao, and J. Pokorny, “Threshold units: a correct metric for reaction time?” Vision Res. 47, 608–611 (2007). [CrossRef] [PubMed]
  57. V. C. Smith and J. Pokorny, “The design and use of a cone chromaticity space,” Color Res. Appl. 21, 375–383 (1996). [CrossRef]
  58. N. V. S. Graham, Visual Pattern Analyzers (Oxford University, 1989). [CrossRef]
  59. V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, and A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221(1992). [PubMed]
  60. J. Kremers, T. Usui, H. P. Scholl, and L. T. Sharpe, “Cone signal contributions to electroretinograms in dichromats and trichromats,” Invest. Ophthalmol. Vis. Sci. 40, 920–930(1999). [PubMed]
  61. N. Benimoff, S. Schneider, and D. C. Hood, “Interactions between rod and cone channels above threshold: a test of various models,” Vis. Res. 22, 1133–1140 (1982). [CrossRef] [PubMed]
  62. B. Drum, “Summation of rod and cone responses at absolute threshold,” Vis. Res. 22, 823–826 (1982). [CrossRef] [PubMed]
  63. S. L. Guth, N. J. Donley, and R. T. Marrocco, “On luminance additivity and related topics,” Vis. Res. 9, 537–575 (1969). [CrossRef] [PubMed]
  64. M. W. Levine and L. J. Frishman, “Interactions between rod and cone channels: a model that includes inhibition,” Vis. Res. 24, 513–516 (1984). [CrossRef] [PubMed]
  65. R. W. Massof and S. J. Starr, “Vector magnitude operation in color vision models: derivation from signal detection theory,” J. Opt. Soc. Am. 70, 870–872 (1980). [CrossRef] [PubMed]
  66. S. L. Buck and R. Knight, “Test-additivity experiments: different procedures, different results,” J. Opt. Soc. Am. A 8, 696–698(1991). [CrossRef] [PubMed]
  67. S. L. Buck and R. Knight, “Partial additivity of rod signals with M- and L-cone signals in increment detection,” Vis. Res. 34, 2537–2545 (1994). [CrossRef] [PubMed]
  68. H. Sun, J. Pokorny, and V. C. Smith, “Control of the modulation of human photoreceptors,” Color Res. Appl. 26, S69–S75 (2001). [CrossRef]
  69. S. L. Buck, “Rod-cone interactions in human vision,” in The Visual Neurosciences, L.M.Chalupa and J.S.Werner, eds. (MIT, 2004), pp. 863–878.
  70. R. Knight, S. L. Buck, and M. Pereverzeva, “Stimulus size affects rod influence on tritan chromatic discrimination,” Color Res. Appl. 26, S65–S68 (2001). [CrossRef]
  71. E. J. Chichilnisky and D. A. Baylor, “Receptive-field microstructure of blue-yellow ganglion cells in primate retina,” Nat. Neurosci. 2, 889–893 (1999). [CrossRef] [PubMed]
  72. D. Cao, A. J. Zele, and J. Pokorny, “Linking impulse response functions to reaction time: rod and cone reaction time data and a computational model,” Vis. Res. 47, 1060–1074 (2007). [CrossRef] [PubMed]
  73. H. E. Smithson and J. D. Mollon, “Is the S-opponent chromatic sub-system sluggish?” Vis. Res. 44, 2919–2929 (2004). [CrossRef] [PubMed]
  74. D. J. McKeefry, N. R. Parry, and I. J. Murray, “Simple reaction times in color space: the influence of chromaticity, contrast, and cone opponency,” Invest. Ophthalmol. Visual Sci. 44, 2267–2276 (2003). [CrossRef]
  75. N. P. Cottaris and R. L. De Valois, “Temporal dynamics of chromatic tuning in macaque primary visual cortex,” Nature 395, 896–900 (1998). [CrossRef] [PubMed]
  76. C. Ripamonti, W. L. Woo, E. Crowther, and A. Stockman, “The S-cone contribution to luminance depends on the M- and L-cone adaptation levels: silent surrounds?” J. Vis. 9, 10 (2009). [CrossRef] [PubMed]
  77. K. Shinomori and J. S. Werner, “The impulse response of S-cone pathways in detection of increments and decrements,” Vis. Neurosci. 25, 341–347 (2008). [CrossRef] [PubMed]
  78. P. Sterling, “Deciphering the retina’s wiring diagram,” Nat. Neurosci. 2, 851–853 (1999). [CrossRef] [PubMed]
  79. D. M. Dacey, B. B. Lee, D. K. Stafford, J. Pokorny, and V. C. Smith, “Horizontal cells of the primate retina: cone specificity without spectral opponency,” Science 271, 656–659(1996). [CrossRef] [PubMed]

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