Abstract

This study examined the detectability of flicker for small long-wavelength foveal test stimuli centered within larger long-wavelength surround stimuli. Flicker visibility was evaluated as a function of surround and test illuminance and as a function of test wavelength, of the time elapsed following test or surround onset, and of surround dimensions. Consistent with prior flicker threshold-versus-illuminance results [ Vision Res. 26, 917 ( 1986)], flicker threshold decreased abruptly once the surround illuminance became sufficiently great. However, as test illuminance was increased above flicker threshold, flicker again vanished. Flicker reappeared at still higher test illuminances, as middle-wavelength-sensitive (M-) cone-mediated flicker threshold was exceeded. Meanwhile, the time required for the surround to render flicker visible increased at a rapidly accelerating rate with decreasing surround illuminance; it increased at a more sporadic rate with increasing test illuminance. At bright enough surround illuminances, flicker did not vanish with increasing test illuminance. These and other results are compatible with a framework derived from previous dark-adaptation data8 [ Vision Res. 32, 1975 ( 1992)]. In that framework the test stimulus itself induces losses of flicker sensitivity by sufficiently perturbing retinal response during states or stages of adaptation that fail to cause spectrally antagonistic processes to redress that perturbation adequately. The relevant adaptation processes, which can require minutes, involve an adaptation pool that includes (and is affected by) the test stimulus.

© 1994 Optical Society of America

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  1. A. Eisner, D. I. A. MacLeod, “Flicker photometric study of chromatic adaptation: selective suppression of cone inputs by colored backgrounds,”J. Opt. Soc. Am. 71, 705–718 (1981).
    [Crossref] [PubMed]
  2. C. W. Tyler, R. D. Hamer, “Analysis of visual modulation sensitivity. IV Validity of the Ferry–Porter law,” J. Opt. Soc. Am. A 7, 743–758 (1990).
    [Crossref] [PubMed]
  3. R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1979).
  4. T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
    [Crossref] [PubMed]
  5. K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
    [Crossref]
  6. N. J. Coletta, A. J. Adams, “Rod–cone interactions in flicker detection,” Vision Res. 24, 1330–1340 (1984).
    [Crossref]
  7. A. Eisner, “Losses of foveal flicker sensitivity during dark adaptation following extended bleaches,” Vision Res. 29, 1401–1423 (1989).
    [Crossref] [PubMed]
  8. A. Eisner, “Losses of flicker sensitivity during dark adaptation: effects of test size and wavelength,” Vision Res. 32, 1975–1986 (1992).
    [Crossref] [PubMed]
  9. A. Eisner, J. R. Samples, “Profound reductions of flicker sensitivity in the elderly: can glaucoma involve the retina distal to ganglion cells?” Appl. Opt. 30, 2121–2135 (1991).
    [Crossref] [PubMed]
  10. N. J. Coletta, A. J. Adams, “Spatial extent of rod–cone interactions for flicker detection,” Vision Res. 26, 917–925 (1986).
    [Crossref]
  11. L. O. Harvey, “Flicker sensitivity and apparent brightness as a function of surround illuminance,”J. Opt. Soc. Am. 60, 860–864 (1970).
    [Crossref]
  12. C. Berger, “Illumination of surrounding field and flicker fusion frequency with foveal images of different sizes,” Acta Physiol. Scand. 30, 161–170 (1954).
    [Crossref] [PubMed]
  13. Subjects dark adapted monocularly rather than binocularly in order to delay any effects of night myopia, which during pilot studies was sometimes observed to cause stimulus defocus toward the end of testing sessions.
  14. I. Powell, “Lenses for correcting chromatic aberration of the eye,” Appl. Opt. 20, 4152–4155 (1981).
    [Crossref] [PubMed]
  15. Four additional subjects had been tested previously with a manual version of the flicker-tvi paradigm. For two of these subjects the threshold for first seeing flicker decreased very steeply at surround illuminances of ~1.0 log Td; for the other two subjects threshold decreased very steeply at ~2.0 log Td. In addition, for subject DL the threshold for first seeing flicker occasionally decreased very steeply at surround illuminances as dim as 1.3 log Td.
  16. For subject TQN the second flicker threshold was usually approximately invariant with surround illuminance. However, for subject DL it only sometimes was, and for subject RH it rarely was. For a fourth subject, tested manually, the second flicker threshold was quite invariant with surround illuminance even though the steep decrease of flicker threshold occurred at ~1.0 log Td.
  17. Flicker threshold sometimes increased by several tenths of a log unit across the first several surround illuminances tested. Control experiments for two subjects (TQN and DL) demonstrated that this effect could occur over time even when the surround remained occluded.
  18. R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
    [Crossref] [PubMed]
  19. L. M. Hurvich, Color Vision (Sinauer, Sunderland, Mass., 1981).
  20. D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
    [Crossref] [PubMed]
  21. V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
    [PubMed]
  22. V. C. Smith, J. Pokorny, appendix, part III, in R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1975), p. 404.
  23. C. M. Cicerone, J. L. Nerger, “The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
    [Crossref] [PubMed]
  24. It is possible that the data at 1.55 log Td in Fig. 4b did not lie to the left of the steep flicker-tvi slope as portrayed. However, delays that exceeded 4 min were recorded at other (noisier or incomplete) testing sessions at the surround illuminance immediately to the left of the steep flicker-tvi slope, as measured in the usual 0.1-log-unit increments.
  25. Data from control experiments for all three subjects imply that flicker did not require a long time to become visible at relatively dim surrounds only because those surrounds were tested last.
  26. M. M. Hayhoe, “Spatial interactions and models of adaptation,” Vision Res. 30, 957–965 (1990).
    [Crossref] [PubMed]
  27. Control experiments for subjects TQN and DL demonstrated that appreciable delays existed at surround illuminances to the left of the steep flicker-tvi slope even after allowance for the possibility that the steep flicker-tvi slope shifted to dimmer surround illuminances because of the surround-onset paradigm itself.
  28. For instance, at one session the times that were required following surround-onset for the flicker to become visible at surround illuminances of 1.8, 1.9, and 2.0 log Td were 42 and 47, 49 and 53, and 30 and 41 s, respectively. The dimmest surround illuminance for which tvi flicker vanished with increasing test illuminance with 1.8 log Td. Data obtained at another testing session were quite similar.
  29. All attempts at stimulus manipulation failed to make subject DL’s functions look like those of subject TQN, which were reliably monotonic. The following parameters were manipulated: test size, test on and off durations, test wavelength, intertrial interval, and surround illuminance.
  30. Control experiments for subjects TQN and RH demonstrated that appreciable delays existed at test illuminances above the vanishing threshold even after allowance for the possibility that the vanishing threshold shifted to higher surround illuminances because of the surround-onset paradigm itself.
  31. Data obtained again at the brighter surround illuminance imply that flicker did not oscillate at the dimmer surround illuminance because of stimulus-order effects.
  32. Flicker remained invisible at four of four sessions for subject TQN, at two of four sessions for subject DL, and at one of two sessions for subject RH. At a test illuminance of 3.0 log Td and a surround illuminance of 2.5 log Td, flicker was visible throughout the entire 5-min period for all three subjects.
  33. J. M. Loomis, “Complementary afterimages and the unequal adapting effects of steady and flickering lights,”J. Opt. Soc. Am. 68, 411–415 (1978).
    [Crossref] [PubMed]
  34. J. M. Loomis, “Transient tritanopia: failure of time-intensity reciprocity in adaptation to longwave light,” Vision Res. 20, 837–846 (1980).
    [Crossref] [PubMed]
  35. There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
    [Crossref] [PubMed]
  36. R. W. Rodieck, “Which cells code for color?” in A. Valberg, B. B. Lee, eds., From Pigments to Perception: Advances in Understanding Visual Processes (Plenum, New York, 1991), pp. 83–91.
    [Crossref]
  37. C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
    [Crossref] [PubMed]
  38. Subject DL was not tested for this experiment.
  39. Any quantitative differences might also have depended on the precise choice of test illuminance or on other sources of between-session variance.
  40. Data from subject TQN demonstrate that the time required for 18′-i.d., 1°-o.d. surrounds to render flicker visible did not increase with surround illuminances up to 3.0 log Td after having first decreased at dimmer surround illuminances (such as those elicited for Fig. 4).
  41. A. G. Shapiro, Q. Zaidi, “The effects of prolonged temporal modulation on the differential response of color mechanisms,” Vision Res. 32, 2065–2075 (1992).
    [Crossref] [PubMed]
  42. W. S. Geisler, “Evidence for the equivalent-background hypothesis in cones,” Vision Res. 19, 799–805 (1979).
    [Crossref] [PubMed]
  43. M. M. Hayhoe, B. Chen, “Temporal modulation sensitivity in cone dark adaptation,” Vision Res. 26, 1715–1725 (1986).
    [Crossref] [PubMed]
  44. D. R. Williams, D. I. A. MacLeod, M. M. Hayhoe, “Punctate sensitivity of the blue-sensitive mechanism,” Vision Res. 21, 1357–1375 (1981).
    [Crossref] [PubMed]
  45. It is not likely that M-cone response to the test stimulus enhances flicker sensitivity by adapting the visual system, since if field spectral sensitivity of flicker enhancement is spectrally opponent [as has been demonstrated in the parafovea (N. J. Coletta, A. J. Adams, “Adaptation of a color-opponent mechanism increases parafoveal sensitivity to luminance flicker,” Vision Res. 26, 1241–1248 (1986))], then an extra M-cone contribution would make the surround less effective rather than more effective.
    [Crossref] [PubMed]
  46. Of course, if M-cone response to a flickering test stimulus is responsible for the shift of the steep flicker-tvi slope to dimmer surround illuminances, it is possible that there is a physiologic M-cone flicker signal at the first (i.e., lower) flicker threshold. For subject TQN the detection threshold at 656 nm was ~0.6 log unit below the corresponding flicker threshold at the dimmest surround illuminance (1.8 log Td) for which there were two flicker thresholds. At a surround illuminance of 0.0 log Td, the detection threshold was ~1.0 log unit less than it was at 1.8 log Td.
  47. R. D. Hamer, C. W. Tyler, “Analysis of visual modulation sensitivity. V. Faster visual response for G- than for R-cone pathway,” J. Opt. Soc. Am. A 9, 1889–1904 (1992).
    [Crossref] [PubMed]
  48. However, because surround illuminance was varied in 0.5-log-unit increments in that study,10 any such effect could easily have been missed. In addition, the estimates of the extent of the lateral cone–cone interactions appeared to differ slightly, depending on whether their extent was measured with use of backgrounds or with use of surrounds.
  49. N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
    [Crossref]
  50. R. Pflug, R. Nelson, “Background enhancement of cone signals in cat horizontal cells,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 240 (1987).
  51. R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
    [PubMed]
  52. M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
    [PubMed]
  53. T. E. Frumkes, S. M. Wu, “Independent influences on cone-mediated responses to light onset and offset in distal retinal neurons,”J. Neurophysiol. 64, 1043–1054 (1990).
    [PubMed]
  54. Experiments that used subjects who have disorders of outer-to-inner-retinal transmission indicate that the adaptation pool could be entirely outer retinal [K. R. Alexander, G. A. Fishman, “A rod–cone interaction in flicker perimetry: evidence for a distal retinal locus,” Doc. Ophthalmol. 60, 3–36 (1985); V. C. Greenstein, D. C. Hood, I. M. Siegel, R. E. Carr, “A possible use of rod–cone interaction in congenital stationary nightblindness,” Clin. Vis. Sci. 3, 69–74 (1988)]. However, these results were rod–cone rather than cone–cone interactions. Furthermore, it is possible that the eyes of those subjects had altered outer-retinal circuitry.
    [Crossref] [PubMed]
  55. M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
    [Crossref] [PubMed]
  56. E. N. Pugh, J. D. Mollon, “A theory of the pi-1 and pi-3 color mechanisms of Stiles,” Vision Res. 19, 293–312 (1979).
    [Crossref]
  57. G. B. Arden, T. E. Frumkes, “Stimulation of rods can increase cone flicker ERG’s in man,” Vision Res. 26, 711–721 (1986).
    [Crossref]
  58. G. B. Arden, C. R. Hogg, “Rod–cone interactions and analysis of retinal disease,” Br. J. Ophthalmol. 69, 404–415 (1985).
    [Crossref] [PubMed]
  59. N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
    [Crossref] [PubMed]
  60. N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
    [PubMed]
  61. N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
    [Crossref] [PubMed]
  62. However, in contrast to those in the present study, the results of Peachey et al.59 were obtained with achromatic test stimuli [presented as ganzfeld “strobe flashes of less than 1-ms duration and of a constant luminance of 0.8 log cd-s/m2” (~2.5 log Td-s) in 31.1-Hz flicker trains for 1.3 s every minute] (p. 146). Nevertheless, the results did appear to be more pronounced for the color-normal subject than for the deuteranomalous subject. In the introduction to their paper, Peachey et al. suggested (p. 145) that the flicker ERG results were “unlikely to be related to the suppression of cone-mediated flicker sensitivity that occurs in the dark… [because] light adaptation … reduces the amplitude of flicker ERG’s when near-threshold stimuli are used.” However, if the flicker-threshold levels in the present study correspond to ERG flicker responses that are sufficiently suprathreshold, then progressively longer times could be required for a suitable electrophysiological criterion to be attained with decreasing adapting field illuminance, just as in the psychophysical experiments.
  63. G. Lange, T. E. Frumkes, “Influence of rod adaptation upon cone responses to light offset in humans. II. Results in an observer with exaggerated suppressive rod–cone interaction,” Vis. Neurosci. 8, 91–95 (1992).
    [Crossref] [PubMed]
  64. N. Graham, D. C. Hood, “Modeling the dynamics of light adaptation: the merging of two traditions,” Vision Res. 32, 1373–1393 (1992).
    [Crossref] [PubMed]
  65. G. Dagnelie, “Temporal impulse responses from flicker sensitivities: practical considerations,” J. Opt. Soc. Am. A 9, 659–672 (1992).
    [Crossref] [PubMed]
  66. C. W. Tyler, “Psychophysical derivation of the impulse response through generation of ultrabrief responses: complex inverse estimation without minimum-phase assumptions,” J. Opt. Soc. Am. A 9, 1025–1040 (1992).
    [Crossref] [PubMed]
  67. R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
    [Crossref] [PubMed]

1992 (14)

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

A. Eisner, “Losses of flicker sensitivity during dark adaptation: effects of test size and wavelength,” Vision Res. 32, 1975–1986 (1992).
[Crossref] [PubMed]

D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
[Crossref] [PubMed]

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

A. G. Shapiro, Q. Zaidi, “The effects of prolonged temporal modulation on the differential response of color mechanisms,” Vision Res. 32, 2065–2075 (1992).
[Crossref] [PubMed]

R. D. Hamer, C. W. Tyler, “Analysis of visual modulation sensitivity. V. Faster visual response for G- than for R-cone pathway,” J. Opt. Soc. Am. A 9, 1889–1904 (1992).
[Crossref] [PubMed]

M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

G. Lange, T. E. Frumkes, “Influence of rod adaptation upon cone responses to light offset in humans. II. Results in an observer with exaggerated suppressive rod–cone interaction,” Vis. Neurosci. 8, 91–95 (1992).
[Crossref] [PubMed]

N. Graham, D. C. Hood, “Modeling the dynamics of light adaptation: the merging of two traditions,” Vision Res. 32, 1373–1393 (1992).
[Crossref] [PubMed]

G. Dagnelie, “Temporal impulse responses from flicker sensitivities: practical considerations,” J. Opt. Soc. Am. A 9, 659–672 (1992).
[Crossref] [PubMed]

C. W. Tyler, “Psychophysical derivation of the impulse response through generation of ultrabrief responses: complex inverse estimation without minimum-phase assumptions,” J. Opt. Soc. Am. A 9, 1025–1040 (1992).
[Crossref] [PubMed]

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

1991 (4)

N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
[PubMed]

M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
[PubMed]

There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
[Crossref] [PubMed]

A. Eisner, J. R. Samples, “Profound reductions of flicker sensitivity in the elderly: can glaucoma involve the retina distal to ganglion cells?” Appl. Opt. 30, 2121–2135 (1991).
[Crossref] [PubMed]

1990 (6)

C. W. Tyler, R. D. Hamer, “Analysis of visual modulation sensitivity. IV Validity of the Ferry–Porter law,” J. Opt. Soc. Am. A 7, 743–758 (1990).
[Crossref] [PubMed]

M. M. Hayhoe, “Spatial interactions and models of adaptation,” Vision Res. 30, 957–965 (1990).
[Crossref] [PubMed]

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
[Crossref]

T. E. Frumkes, S. M. Wu, “Independent influences on cone-mediated responses to light onset and offset in distal retinal neurons,”J. Neurophysiol. 64, 1043–1054 (1990).
[PubMed]

R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
[PubMed]

1989 (3)

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

C. M. Cicerone, J. L. Nerger, “The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
[Crossref] [PubMed]

A. Eisner, “Losses of foveal flicker sensitivity during dark adaptation following extended bleaches,” Vision Res. 29, 1401–1423 (1989).
[Crossref] [PubMed]

1988 (1)

K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
[Crossref]

1987 (1)

R. Pflug, R. Nelson, “Background enhancement of cone signals in cat horizontal cells,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 240 (1987).

1986 (4)

M. M. Hayhoe, B. Chen, “Temporal modulation sensitivity in cone dark adaptation,” Vision Res. 26, 1715–1725 (1986).
[Crossref] [PubMed]

It is not likely that M-cone response to the test stimulus enhances flicker sensitivity by adapting the visual system, since if field spectral sensitivity of flicker enhancement is spectrally opponent [as has been demonstrated in the parafovea (N. J. Coletta, A. J. Adams, “Adaptation of a color-opponent mechanism increases parafoveal sensitivity to luminance flicker,” Vision Res. 26, 1241–1248 (1986))], then an extra M-cone contribution would make the surround less effective rather than more effective.
[Crossref] [PubMed]

G. B. Arden, T. E. Frumkes, “Stimulation of rods can increase cone flicker ERG’s in man,” Vision Res. 26, 711–721 (1986).
[Crossref]

N. J. Coletta, A. J. Adams, “Spatial extent of rod–cone interactions for flicker detection,” Vision Res. 26, 917–925 (1986).
[Crossref]

1985 (2)

G. B. Arden, C. R. Hogg, “Rod–cone interactions and analysis of retinal disease,” Br. J. Ophthalmol. 69, 404–415 (1985).
[Crossref] [PubMed]

Experiments that used subjects who have disorders of outer-to-inner-retinal transmission indicate that the adaptation pool could be entirely outer retinal [K. R. Alexander, G. A. Fishman, “A rod–cone interaction in flicker perimetry: evidence for a distal retinal locus,” Doc. Ophthalmol. 60, 3–36 (1985); V. C. Greenstein, D. C. Hood, I. M. Siegel, R. E. Carr, “A possible use of rod–cone interaction in congenital stationary nightblindness,” Clin. Vis. Sci. 3, 69–74 (1988)]. However, these results were rod–cone rather than cone–cone interactions. Furthermore, it is possible that the eyes of those subjects had altered outer-retinal circuitry.
[Crossref] [PubMed]

1984 (1)

N. J. Coletta, A. J. Adams, “Rod–cone interactions in flicker detection,” Vision Res. 24, 1330–1340 (1984).
[Crossref]

1981 (3)

1980 (1)

J. M. Loomis, “Transient tritanopia: failure of time-intensity reciprocity in adaptation to longwave light,” Vision Res. 20, 837–846 (1980).
[Crossref] [PubMed]

1979 (2)

W. S. Geisler, “Evidence for the equivalent-background hypothesis in cones,” Vision Res. 19, 799–805 (1979).
[Crossref] [PubMed]

E. N. Pugh, J. D. Mollon, “A theory of the pi-1 and pi-3 color mechanisms of Stiles,” Vision Res. 19, 293–312 (1979).
[Crossref]

1978 (1)

1970 (1)

1954 (1)

C. Berger, “Illumination of surrounding field and flicker fusion frequency with foveal images of different sizes,” Acta Physiol. Scand. 30, 161–170 (1954).
[Crossref] [PubMed]

Adams, A. J.

N. J. Coletta, A. J. Adams, “Spatial extent of rod–cone interactions for flicker detection,” Vision Res. 26, 917–925 (1986).
[Crossref]

It is not likely that M-cone response to the test stimulus enhances flicker sensitivity by adapting the visual system, since if field spectral sensitivity of flicker enhancement is spectrally opponent [as has been demonstrated in the parafovea (N. J. Coletta, A. J. Adams, “Adaptation of a color-opponent mechanism increases parafoveal sensitivity to luminance flicker,” Vision Res. 26, 1241–1248 (1986))], then an extra M-cone contribution would make the surround less effective rather than more effective.
[Crossref] [PubMed]

N. J. Coletta, A. J. Adams, “Rod–cone interactions in flicker detection,” Vision Res. 24, 1330–1340 (1984).
[Crossref]

Alexander, K. R.

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
[PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
[Crossref]

K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
[Crossref]

Experiments that used subjects who have disorders of outer-to-inner-retinal transmission indicate that the adaptation pool could be entirely outer retinal [K. R. Alexander, G. A. Fishman, “A rod–cone interaction in flicker perimetry: evidence for a distal retinal locus,” Doc. Ophthalmol. 60, 3–36 (1985); V. C. Greenstein, D. C. Hood, I. M. Siegel, R. E. Carr, “A possible use of rod–cone interaction in congenital stationary nightblindness,” Clin. Vis. Sci. 3, 69–74 (1988)]. However, these results were rod–cone rather than cone–cone interactions. Furthermore, it is possible that the eyes of those subjects had altered outer-retinal circuitry.
[Crossref] [PubMed]

Arakawa, K.

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

Arden, G. B.

M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
[PubMed]

G. B. Arden, T. E. Frumkes, “Stimulation of rods can increase cone flicker ERG’s in man,” Vision Res. 26, 711–721 (1986).
[Crossref]

G. B. Arden, C. R. Hogg, “Rod–cone interactions and analysis of retinal disease,” Br. J. Ophthalmol. 69, 404–415 (1985).
[Crossref] [PubMed]

Baer, S. M.

R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
[PubMed]

Beczkowska, I.

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

Berger, C.

C. Berger, “Illumination of surrounding field and flicker fusion frequency with foveal images of different sizes,” Acta Physiol. Scand. 30, 161–170 (1954).
[Crossref] [PubMed]

Bowen, R. W.

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

Boynton, R. M.

R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1979).

Calkins, D. J.

D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
[Crossref] [PubMed]

Chen, B.

M. M. Hayhoe, B. Chen, “Temporal modulation sensitivity in cone dark adaptation,” Vision Res. 26, 1715–1725 (1986).
[Crossref] [PubMed]

Cicerone, C. M.

C. M. Cicerone, J. L. Nerger, “The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
[Crossref] [PubMed]

Coletta, N. J.

N. J. Coletta, A. J. Adams, “Spatial extent of rod–cone interactions for flicker detection,” Vision Res. 26, 917–925 (1986).
[Crossref]

It is not likely that M-cone response to the test stimulus enhances flicker sensitivity by adapting the visual system, since if field spectral sensitivity of flicker enhancement is spectrally opponent [as has been demonstrated in the parafovea (N. J. Coletta, A. J. Adams, “Adaptation of a color-opponent mechanism increases parafoveal sensitivity to luminance flicker,” Vision Res. 26, 1241–1248 (1986))], then an extra M-cone contribution would make the surround less effective rather than more effective.
[Crossref] [PubMed]

N. J. Coletta, A. J. Adams, “Rod–cone interactions in flicker detection,” Vision Res. 24, 1330–1340 (1984).
[Crossref]

Curcio, C. A.

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

Dagnelie, G.

Denny, N.

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

Derlacki, D. J.

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
[Crossref]

K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
[Crossref]

Eisner, A.

Eysteinsson, T.

M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
[PubMed]

Fishman, G. A.

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
[PubMed]

K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
[Crossref]

Experiments that used subjects who have disorders of outer-to-inner-retinal transmission indicate that the adaptation pool could be entirely outer retinal [K. R. Alexander, G. A. Fishman, “A rod–cone interaction in flicker perimetry: evidence for a distal retinal locus,” Doc. Ophthalmol. 60, 3–36 (1985); V. C. Greenstein, D. C. Hood, I. M. Siegel, R. E. Carr, “A possible use of rod–cone interaction in congenital stationary nightblindness,” Clin. Vis. Sci. 3, 69–74 (1988)]. However, these results were rod–cone rather than cone–cone interactions. Furthermore, it is possible that the eyes of those subjects had altered outer-retinal circuitry.
[Crossref] [PubMed]

Fowler, M. A.

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

Frumkes, T. E.

G. Lange, T. E. Frumkes, “Influence of rod adaptation upon cone responses to light offset in humans. II. Results in an observer with exaggerated suppressive rod–cone interaction,” Vis. Neurosci. 8, 91–95 (1992).
[Crossref] [PubMed]

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

T. E. Frumkes, S. M. Wu, “Independent influences on cone-mediated responses to light onset and offset in distal retinal neurons,”J. Neurophysiol. 64, 1043–1054 (1990).
[PubMed]

G. B. Arden, T. E. Frumkes, “Stimulation of rods can increase cone flicker ERG’s in man,” Vision Res. 26, 711–721 (1986).
[Crossref]

Geisler, W. S.

W. S. Geisler, “Evidence for the equivalent-background hypothesis in cones,” Vision Res. 19, 799–805 (1979).
[Crossref] [PubMed]

Graham, N.

N. Graham, D. C. Hood, “Modeling the dynamics of light adaptation: the merging of two traditions,” Vision Res. 32, 1373–1393 (1992).
[Crossref] [PubMed]

Hamer, R. D.

Harvey, L. O.

Hayhoe, M. M.

M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
[Crossref] [PubMed]

M. M. Hayhoe, “Spatial interactions and models of adaptation,” Vision Res. 30, 957–965 (1990).
[Crossref] [PubMed]

M. M. Hayhoe, B. Chen, “Temporal modulation sensitivity in cone dark adaptation,” Vision Res. 26, 1715–1725 (1986).
[Crossref] [PubMed]

D. R. Williams, D. I. A. MacLeod, M. M. Hayhoe, “Punctate sensitivity of the blue-sensitive mechanism,” Vision Res. 21, 1357–1375 (1981).
[Crossref] [PubMed]

Hendrickson, A. E.

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

Hogg, C. R.

G. B. Arden, C. R. Hogg, “Rod–cone interactions and analysis of retinal disease,” Br. J. Ophthalmol. 69, 404–415 (1985).
[Crossref] [PubMed]

Hood, D. C.

N. Graham, D. C. Hood, “Modeling the dynamics of light adaptation: the merging of two traditions,” Vision Res. 32, 1373–1393 (1992).
[Crossref] [PubMed]

Horiguchi, M.

M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
[PubMed]

Hurvich, L. M.

L. M. Hurvich, Color Vision (Sinauer, Sunderland, Mass., 1981).

Kalina, R. E.

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

Koshel, R. J.

M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
[Crossref] [PubMed]

Lange, G.

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

G. Lange, T. E. Frumkes, “Influence of rod adaptation upon cone responses to light offset in humans. II. Results in an observer with exaggerated suppressive rod–cone interaction,” Vis. Neurosci. 8, 91–95 (1992).
[Crossref] [PubMed]

Lee, B. B.

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

Levin, M. E.

M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
[Crossref] [PubMed]

Loomis, J. M.

J. M. Loomis, “Transient tritanopia: failure of time-intensity reciprocity in adaptation to longwave light,” Vision Res. 20, 837–846 (1980).
[Crossref] [PubMed]

J. M. Loomis, “Complementary afterimages and the unequal adapting effects of steady and flickering lights,”J. Opt. Soc. Am. 68, 411–415 (1978).
[Crossref] [PubMed]

MacLeod, D. I. A.

Marchese, A. L.

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

Martin, P. R.

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

Mollon, J. D.

E. N. Pugh, J. D. Mollon, “A theory of the pi-1 and pi-3 color mechanisms of Stiles,” Vision Res. 19, 293–312 (1979).
[Crossref]

Nelson, R.

R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
[PubMed]

R. Pflug, R. Nelson, “Background enhancement of cone signals in cat horizontal cells,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 240 (1987).

Nerger, J. L.

C. M. Cicerone, J. L. Nerger, “The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
[Crossref] [PubMed]

Peachey, N. S.

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
[PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
[Crossref]

Petrig, B. L.

There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
[Crossref] [PubMed]

Pflug, R.

R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
[PubMed]

R. Pflug, R. Nelson, “Background enhancement of cone signals in cat horizontal cells,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 240 (1987).

Pokorny, J.

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

V. C. Smith, J. Pokorny, appendix, part III, in R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1975), p. 404.

Powell, I.

Pugh, E. N.

D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
[Crossref] [PubMed]

E. N. Pugh, J. D. Mollon, “A theory of the pi-1 and pi-3 color mechanisms of Stiles,” Vision Res. 19, 293–312 (1979).
[Crossref]

Riva, C. E.

There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
[Crossref] [PubMed]

Rodieck, R. W.

R. W. Rodieck, “Which cells code for color?” in A. Valberg, B. B. Lee, eds., From Pigments to Perception: Advances in Understanding Visual Processes (Plenum, New York, 1991), pp. 83–91.
[Crossref]

Samples, J. R.

Shapiro, A. G.

A. G. Shapiro, Q. Zaidi, “The effects of prolonged temporal modulation on the differential response of color mechanisms,” Vision Res. 32, 2065–2075 (1992).
[Crossref] [PubMed]

Shevell, S. K.

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

Shonet, R. D.

There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
[Crossref] [PubMed]

Sloan, K. R.

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

Smith, V. C.

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

V. C. Smith, J. Pokorny, appendix, part III, in R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1975), p. 404.

Thornton, J. E.

D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
[Crossref] [PubMed]

Tyler, C. W.

Valberg, A.

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

Vimal, R. L. P.

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

Williams, D. R.

D. R. Williams, D. I. A. MacLeod, M. M. Hayhoe, “Punctate sensitivity of the blue-sensitive mechanism,” Vision Res. 21, 1357–1375 (1981).
[Crossref] [PubMed]

Wu, S. M.

T. E. Frumkes, S. M. Wu, “Independent influences on cone-mediated responses to light onset and offset in distal retinal neurons,”J. Neurophysiol. 64, 1043–1054 (1990).
[PubMed]

Zaidi, Q.

A. G. Shapiro, Q. Zaidi, “The effects of prolonged temporal modulation on the differential response of color mechanisms,” Vision Res. 32, 2065–2075 (1992).
[Crossref] [PubMed]

Acta Physiol. Scand. (1)

C. Berger, “Illumination of surrounding field and flicker fusion frequency with foveal images of different sizes,” Acta Physiol. Scand. 30, 161–170 (1954).
[Crossref] [PubMed]

Appl. Opt. (2)

Br. J. Ophthalmol. (1)

G. B. Arden, C. R. Hogg, “Rod–cone interactions and analysis of retinal disease,” Br. J. Ophthalmol. 69, 404–415 (1985).
[Crossref] [PubMed]

Doc. Ophthalmol. (1)

Experiments that used subjects who have disorders of outer-to-inner-retinal transmission indicate that the adaptation pool could be entirely outer retinal [K. R. Alexander, G. A. Fishman, “A rod–cone interaction in flicker perimetry: evidence for a distal retinal locus,” Doc. Ophthalmol. 60, 3–36 (1985); V. C. Greenstein, D. C. Hood, I. M. Siegel, R. E. Carr, “A possible use of rod–cone interaction in congenital stationary nightblindness,” Clin. Vis. Sci. 3, 69–74 (1988)]. However, these results were rod–cone rather than cone–cone interactions. Furthermore, it is possible that the eyes of those subjects had altered outer-retinal circuitry.
[Crossref] [PubMed]

Invest. Ophthalmol. Vis. Sci. (2)

M. Horiguchi, T. Eysteinsson, G. B. Arden, “Temporal and spatial properties of suppressive rod–cone interaction,” Invest. Ophthalmol. Vis. Sci. 32, 575–581 (1991).
[PubMed]

N. S. Peachey, K. R. Alexander, G. A. Fishman, “Visual adaptation and the cone flicker electroretinogram,” Invest. Ophthalmol. Vis. Sci. 32, 1517–1522 (1991).
[PubMed]

Invest. Ophthalmol. Vis. Sci. Suppl. (1)

R. Pflug, R. Nelson, “Background enhancement of cone signals in cat horizontal cells,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 240 (1987).

J. Comp. Neurol. (1)

C. A. Curcio, K. R. Sloan, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,”J. Comp. Neurol. 292, 497–523 (1990).
[Crossref] [PubMed]

J. Neurophysiol. (2)

R. Nelson, R. Pflug, S. M. Baer, “Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties,”J. Neurophysiol. 64, 326–340 (1990).
[PubMed]

T. E. Frumkes, S. M. Wu, “Independent influences on cone-mediated responses to light onset and offset in distal retinal neurons,”J. Neurophysiol. 64, 1043–1054 (1990).
[PubMed]

J. Opt. Soc. Am. (3)

J. Opt. Soc. Am. A (4)

J. Physiol. (1)

V. C. Smith, B. B. Lee, J. Pokorny, P. R. Martin, A. Valberg, “Responses of macaque ganglion cells to the relative phase of heterochromatically modulated lights,” J. Physiol. 458, 191–221 (1992). As Smith et al. note, their results for both on and off cells can be explained either by an excitatory +L–M mechanism that adds to center activity or by a +M–L mechanism that inhibits center activity.
[PubMed]

Neurosci. Lett. (1)

There is even a remote possibility that oscillations could involve flicker-dependent alterations of ocular blood flow. [C. E. Riva, R. D. Shonet, B. L. Petrig, “Flicker evoked increase in optic nerve head blood flow in anesthetized cats,” Neurosci. Lett. 128, 291–296 (1991).]
[Crossref] [PubMed]

Vis. Neurosci. (3)

G. Lange, T. E. Frumkes, “Influence of rod adaptation upon cone responses to light offset in humans. II. Results in an observer with exaggerated suppressive rod–cone interaction,” Vis. Neurosci. 8, 91–95 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, G. A. Fishman, “Light adaptation, rods and the human flicker ERG,” Vis. Neurosci. 8, 145–150 (1992).
[Crossref] [PubMed]

T. E. Frumkes, G. Lange, N. Denny, I. Beczkowska, “Influence of rod adaptation upon cone responses to light offset in humans. I. Results in normal observers,” Vis. Neurosci. 8, 83–89 (1992).
[Crossref] [PubMed]

Vision Res. (22)

K. R. Alexander, G. A. Fishman, D. J. Derlacki, “Mechanisms of rod–cone interaction: evidence from congenital stationary night blindness,” Vision Res. 28, 575–583 (1988).
[Crossref]

N. J. Coletta, A. J. Adams, “Rod–cone interactions in flicker detection,” Vision Res. 24, 1330–1340 (1984).
[Crossref]

A. Eisner, “Losses of foveal flicker sensitivity during dark adaptation following extended bleaches,” Vision Res. 29, 1401–1423 (1989).
[Crossref] [PubMed]

A. Eisner, “Losses of flicker sensitivity during dark adaptation: effects of test size and wavelength,” Vision Res. 32, 1975–1986 (1992).
[Crossref] [PubMed]

N. J. Coletta, A. J. Adams, “Spatial extent of rod–cone interactions for flicker detection,” Vision Res. 26, 917–925 (1986).
[Crossref]

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[Crossref] [PubMed]

D. J. Calkins, J. E. Thornton, E. N. Pugh, “Monochromatism determined at a long-wavelength/middle-wavelength cone-antagonistic locus,” Vision Res. 32, 2349–2367 (1992).
[Crossref] [PubMed]

M. M. Hayhoe, “Spatial interactions and models of adaptation,” Vision Res. 30, 957–965 (1990).
[Crossref] [PubMed]

J. M. Loomis, “Transient tritanopia: failure of time-intensity reciprocity in adaptation to longwave light,” Vision Res. 20, 837–846 (1980).
[Crossref] [PubMed]

C. M. Cicerone, J. L. Nerger, “The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
[Crossref] [PubMed]

N. S. Peachey, K. Arakawa, K. R. Alexander, A. L. Marchese, “Rapid and slow changes in the human cone electroretinogram during light and dark adaptation,” Vision Res. 32, 2049–2053 (1992).
[Crossref] [PubMed]

N. Graham, D. C. Hood, “Modeling the dynamics of light adaptation: the merging of two traditions,” Vision Res. 32, 1373–1393 (1992).
[Crossref] [PubMed]

R. W. Bowen, J. Pokorny, V. C. Smith, M. A. Fowler, “Sawtooth contrast sensitivity: effects of mean illuminance and low temporal frequencies,” Vision Res. 32, 1239–1247 (1992).
[Crossref] [PubMed]

N. S. Peachey, K. R. Alexander, D. J. Derlacki, “Spatial properties of rod–cone interactions in flicker and hue detection,” Vision Res. 30, 1205–1210 (1990).
[Crossref]

A. G. Shapiro, Q. Zaidi, “The effects of prolonged temporal modulation on the differential response of color mechanisms,” Vision Res. 32, 2065–2075 (1992).
[Crossref] [PubMed]

W. S. Geisler, “Evidence for the equivalent-background hypothesis in cones,” Vision Res. 19, 799–805 (1979).
[Crossref] [PubMed]

M. M. Hayhoe, B. Chen, “Temporal modulation sensitivity in cone dark adaptation,” Vision Res. 26, 1715–1725 (1986).
[Crossref] [PubMed]

D. R. Williams, D. I. A. MacLeod, M. M. Hayhoe, “Punctate sensitivity of the blue-sensitive mechanism,” Vision Res. 21, 1357–1375 (1981).
[Crossref] [PubMed]

It is not likely that M-cone response to the test stimulus enhances flicker sensitivity by adapting the visual system, since if field spectral sensitivity of flicker enhancement is spectrally opponent [as has been demonstrated in the parafovea (N. J. Coletta, A. J. Adams, “Adaptation of a color-opponent mechanism increases parafoveal sensitivity to luminance flicker,” Vision Res. 26, 1241–1248 (1986))], then an extra M-cone contribution would make the surround less effective rather than more effective.
[Crossref] [PubMed]

M. M. Hayhoe, M. E. Levin, R. J. Koshel, “Subtractive processes in light adaptation,” Vision Res. 32, 323–333 (1992).
[Crossref] [PubMed]

E. N. Pugh, J. D. Mollon, “A theory of the pi-1 and pi-3 color mechanisms of Stiles,” Vision Res. 19, 293–312 (1979).
[Crossref]

G. B. Arden, T. E. Frumkes, “Stimulation of rods can increase cone flicker ERG’s in man,” Vision Res. 26, 711–721 (1986).
[Crossref]

Other (22)

Of course, if M-cone response to a flickering test stimulus is responsible for the shift of the steep flicker-tvi slope to dimmer surround illuminances, it is possible that there is a physiologic M-cone flicker signal at the first (i.e., lower) flicker threshold. For subject TQN the detection threshold at 656 nm was ~0.6 log unit below the corresponding flicker threshold at the dimmest surround illuminance (1.8 log Td) for which there were two flicker thresholds. At a surround illuminance of 0.0 log Td, the detection threshold was ~1.0 log unit less than it was at 1.8 log Td.

R. W. Rodieck, “Which cells code for color?” in A. Valberg, B. B. Lee, eds., From Pigments to Perception: Advances in Understanding Visual Processes (Plenum, New York, 1991), pp. 83–91.
[Crossref]

However, because surround illuminance was varied in 0.5-log-unit increments in that study,10 any such effect could easily have been missed. In addition, the estimates of the extent of the lateral cone–cone interactions appeared to differ slightly, depending on whether their extent was measured with use of backgrounds or with use of surrounds.

However, in contrast to those in the present study, the results of Peachey et al.59 were obtained with achromatic test stimuli [presented as ganzfeld “strobe flashes of less than 1-ms duration and of a constant luminance of 0.8 log cd-s/m2” (~2.5 log Td-s) in 31.1-Hz flicker trains for 1.3 s every minute] (p. 146). Nevertheless, the results did appear to be more pronounced for the color-normal subject than for the deuteranomalous subject. In the introduction to their paper, Peachey et al. suggested (p. 145) that the flicker ERG results were “unlikely to be related to the suppression of cone-mediated flicker sensitivity that occurs in the dark… [because] light adaptation … reduces the amplitude of flicker ERG’s when near-threshold stimuli are used.” However, if the flicker-threshold levels in the present study correspond to ERG flicker responses that are sufficiently suprathreshold, then progressively longer times could be required for a suitable electrophysiological criterion to be attained with decreasing adapting field illuminance, just as in the psychophysical experiments.

It is possible that the data at 1.55 log Td in Fig. 4b did not lie to the left of the steep flicker-tvi slope as portrayed. However, delays that exceeded 4 min were recorded at other (noisier or incomplete) testing sessions at the surround illuminance immediately to the left of the steep flicker-tvi slope, as measured in the usual 0.1-log-unit increments.

Data from control experiments for all three subjects imply that flicker did not require a long time to become visible at relatively dim surrounds only because those surrounds were tested last.

Subject DL was not tested for this experiment.

Any quantitative differences might also have depended on the precise choice of test illuminance or on other sources of between-session variance.

Data from subject TQN demonstrate that the time required for 18′-i.d., 1°-o.d. surrounds to render flicker visible did not increase with surround illuminances up to 3.0 log Td after having first decreased at dimmer surround illuminances (such as those elicited for Fig. 4).

Control experiments for subjects TQN and DL demonstrated that appreciable delays existed at surround illuminances to the left of the steep flicker-tvi slope even after allowance for the possibility that the steep flicker-tvi slope shifted to dimmer surround illuminances because of the surround-onset paradigm itself.

For instance, at one session the times that were required following surround-onset for the flicker to become visible at surround illuminances of 1.8, 1.9, and 2.0 log Td were 42 and 47, 49 and 53, and 30 and 41 s, respectively. The dimmest surround illuminance for which tvi flicker vanished with increasing test illuminance with 1.8 log Td. Data obtained at another testing session were quite similar.

All attempts at stimulus manipulation failed to make subject DL’s functions look like those of subject TQN, which were reliably monotonic. The following parameters were manipulated: test size, test on and off durations, test wavelength, intertrial interval, and surround illuminance.

Control experiments for subjects TQN and RH demonstrated that appreciable delays existed at test illuminances above the vanishing threshold even after allowance for the possibility that the vanishing threshold shifted to higher surround illuminances because of the surround-onset paradigm itself.

Data obtained again at the brighter surround illuminance imply that flicker did not oscillate at the dimmer surround illuminance because of stimulus-order effects.

Flicker remained invisible at four of four sessions for subject TQN, at two of four sessions for subject DL, and at one of two sessions for subject RH. At a test illuminance of 3.0 log Td and a surround illuminance of 2.5 log Td, flicker was visible throughout the entire 5-min period for all three subjects.

V. C. Smith, J. Pokorny, appendix, part III, in R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1975), p. 404.

L. M. Hurvich, Color Vision (Sinauer, Sunderland, Mass., 1981).

Subjects dark adapted monocularly rather than binocularly in order to delay any effects of night myopia, which during pilot studies was sometimes observed to cause stimulus defocus toward the end of testing sessions.

Four additional subjects had been tested previously with a manual version of the flicker-tvi paradigm. For two of these subjects the threshold for first seeing flicker decreased very steeply at surround illuminances of ~1.0 log Td; for the other two subjects threshold decreased very steeply at ~2.0 log Td. In addition, for subject DL the threshold for first seeing flicker occasionally decreased very steeply at surround illuminances as dim as 1.3 log Td.

For subject TQN the second flicker threshold was usually approximately invariant with surround illuminance. However, for subject DL it only sometimes was, and for subject RH it rarely was. For a fourth subject, tested manually, the second flicker threshold was quite invariant with surround illuminance even though the steep decrease of flicker threshold occurred at ~1.0 log Td.

Flicker threshold sometimes increased by several tenths of a log unit across the first several surround illuminances tested. Control experiments for two subjects (TQN and DL) demonstrated that this effect could occur over time even when the surround remained occluded.

R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1979).

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Figures (7)

Fig. 1
Fig. 1

a, Flicker-tvi data for 656-nm, 13′-diameter foveal test stimuli centered within 18′-inner-diameter, 1°-outer-diameter, 660-nm surround stimuli. Flicker thresholds (open symbols) and vanishing thresholds (filled symbols) are plotted as functions of surround illuminance. Solid curves connect mean data from two excursions per surround illuminance. Dashed curves connect mean data obtained with no surround to mean data obtained with the dimmest surround. Threshold levels are peak illuminances of the 18 Hz, 100%-modulated sinusoidal test stimuli. Subject: TQN. b, Same as a but for subject RH. c, Same as a but for subject DL.

Fig. 2
Fig. 2

a, Flicker-tvi curves for 620- and 656-nm tests. Other details as in Fig. 1, except that there is only one excursion per surround illuminance per test wavelength. Subject TQN. b, same as a but for subject RH. c, Same as a but for subject DL.

Fig. 3
Fig. 3

a, Flicker spectral sensitivities at two surround illuminances, specified on the graph. Downward- or leftward-pointing triangles represent data collected in descending order of test wavelength; upward- or rightward-pointing triangles represent data collected in ascending order. Upper solid curve, L-cone spectral sensitivity; lower solid curve, M-cone spectral sensitivity; dashed curve, Vλ. Curves are positioned through the data to minimize root-mean-squared deviation from the mean data across test wavelengths. Other details as in Fig. 1. Subject TQN. b, Same as a but for subject RH. c, Same as a but for subject DL.

Fig. 4
Fig. 4

a, Time required for the test to reach flicker threshold after surround onset as a function of surround illuminance. Same stimulus parameters as for Fig. 1. Two traverses of surround illuminance were made, beginning at a surround illuminance of 1.90 log Td and then initially ascending. Upward- and downward-pointing triangles represent ascending and descending orders, respectively, of surround-illuminance variation. The upward-pointing arrow signifies that flicker did not meet flicker-threshold criteria within a 5-min period. The vertical dashed line separates surround illuminances with only a single elevated flicker-tvi threshold from surround illuminances with two flicker-tvi thresholds (i.e., with one vanishing threshold). Test illuminance was 2.32 log Td. At the 1.90-log-Td surround illuminance, the first flicker-tvi threshold was 1.92 log Td and the vanishing threshold was 2.32 log Td. Subject TQN. b, Same as a, except that traverses began at a surround illuminance of 1.70 log Td, which was the second surround illuminance for which flicker was found to vanish. The leftward-pointing triangle represents the last datum of the session. Test illuminance was 2.42 log Td. At the 1.60-log-Td surround illuminance, the first flicker-tvi threshold was 2.22 log Td and the vanishing threshold was 2.62 log Td. Subject RH. c, Same as a, except that traverses began at a surround illuminance of 1.60 log Td. The rightward-pointing triangle represents the last datum of the session. Test illuminance was 2.52 log Td. At the 1.60-log-Td surround illuminance, the first flicker-tvi threshold was 2.32 log Td and the vanishing threshold was 2.62 log Td. Subject DL.

Fig. 5
Fig. 5

a, Time required for the test to reach flicker threshold after surround onset as a function of test illuminance. Two traverses of test illuminance were made, beginning at a test illuminance of 2.10 log Td and then initially ascending. Upward- and downward-pointing triangles represent ascending and descending orders, respectively, of test-illuminance variation. The upward-pointing arrow signifies that flicker did not meet flicker-threshold criteria within a 5-min period. The vertical dashed line separates test illuminances for which flicker vanished in the tvi paradigm from the lower test illuminances for which it did not. The surround illuminance was 1.90 log Td. Subject TQN. b, Same as a, except that traverses began at a test illuminance of 2.50 log Td. The surround illuminance was 1.90 log Td. Subject RH. c, Same as a, except that traverses began at a test illuminance of 2.60 log Td. The surround illuminance was 1.80 log Td. Subject DL.

Fig. 6
Fig. 6

a, Flicker-tvi data for 656-nm, 13′-diameter foveal test stimuli centered within 30′-inner-diameter, 45′-outer-diameter, 660-nm surround stimuli. Other details as in Fig. 1 but for only one excursion per surround illuminance. Subject TQN. b, Same as a but for subject RH. c, Same as a but for subject DL.

Fig. 7
Fig. 7

a, Time required for the text to reach flicker threshold after surround onset as a function of surround illuminance. Same stimulus parameters as for Fig. 6. Same symbols and methodology as for Fig. 4. Traverses began at a surround illuminance of 3.10 log Td. Test illuminance was 2.38 log Td. The first flicker-tvi thresholds at 3.10 log Td were 2.03 and 2.22, and the vanishing thresholds were 2.53 and 2.52 (two excursions). Subject TQN. b, Same as a, except that traverses began at a surround illuminance of 2.70 log Td. Test illuminance was 2.42 log Td. The first flicker-tvi thresholds at 2.70 log Td were 2.42 and 2.12 log Td, and the vanishing thresholds were 2.62 and 2.62 log Td (two excursions). Subject RH.

Tables (1)

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Table 1 Time Required for Flicker to Appear and Disappear following Test Onset after Long-Term Surround Adaptation for Three Subjectsa

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