Abstract

Threshold energy of foveal, 2-msec white test flashes was plotted as a function of their timing with respect to sinusoidal fluctuations of nearly achromatic background light. With frequencies of 3.1–10 Hz, modulation near 100%, and peak retinal illuminance of 1280 td, the resulting masking curves are nonsinusoidal. Threshold maxima precede background maxima, whereas threshold minima coincide with background minima. The phase lead of the threshold maxima increases with frequency, but tends to stabilize at about 90° above 5 Hz. Maximal and minimal threshold energies also increase with frequency, but their ratio is nearly constant between 3.1 and 8 Hz. Steady light added to the 1280-td fluctuation elevates threshold minima, but often lowers maxima, and introduces secondary maxima that accompany decrements of background luminance. Generally, thresholds are higher with fluctuating backgrounds than with peak-equivalent steady backgrounds. The data for 1280-td and 128-td fluctuations confirm that masking is a nonlinear function of modulated background luminance, and suggest the presence of unidirectional rate sensitivity. Of several possible physiological correlates discussed, ganglion-cell discharge patterns are most nearly compatible with the masking curves.

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  1. G. Sperling, Doc. Ophthalmol. 18, 3 (1964).
    [Crossref]
  2. P. Gouras and R. D. Gunkel, Doc. Ophthalmol. 18, 137 (1964).
    [Crossref]
  3. B. Cleland and C. Enroth-Cugell, Acta Physiol. Scand. 68, 365 (1966).
    [Crossref]
  4. W. Rackensperger and O.-J. Grüsser, Experientia 22, 192 (1966).
    [Crossref] [PubMed]
  5. L. H. van der Tweel, Ann. N. Y. Acad. Sci. 89, 829 (1961).
    [Crossref] [PubMed]
  6. L. H. van der Tweel and H. F. E. Verduyn Lunel, EEG Clin. Neurophysiol. 18, 587 (1965).
    [Crossref]
  7. M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
    [Crossref] [PubMed]
  8. D. Kahneman, Psychol. Bull. 70, 404 (1968).
    [Crossref] [PubMed]
  9. G. Sperling, J. Opt. Soc. Am. 55, 541 (1965).
    [Crossref]
  10. R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
    [Crossref]
  11. M. Alpern and W. A. H. Rushton, J. Physiol. (London) 189, 519 (1967).
  12. H. deLange, J. Opt. Soc. Am. 44, 380 (1954).
    [Crossref]
  13. H. deLange, J. Opt. Soc. Am. 48, 777 (1958).
    [Crossref]
  14. G. Sperling and M. M. Sondhi, J. Opt. Soc. Am. 58, 1133 (1968).
    [Crossref] [PubMed]
  15. G. Sperling, J. Opt. Soc. Am. 53, 520 (1963).
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    [Crossref] [PubMed]
  17. D. H. Kelly, Doc. Ophthahlol. 18, 16 (1964).
    [Crossref]
  18. L. Matin, Am: J. Psychol. 77, 650 (1964).
    [Crossref]
  19. G. Westheimer, Vision Res. 6, 669 (1966).
    [Crossref] [PubMed]
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    [Crossref] [PubMed]
  21. R. Granit, Sensory Mechanisms of the Retina (Hafner Publishing Co., N. Y., 1963), Ch. 8, p. 145.
  22. M. G. F. Fuortes and A. L. Hodgkin, J. Physiol. (London) 172, 239 (1964).
  23. W. A. H. Rushton, J. Physiol. (London) 181, 645 (1965).
  24. R. A. Cone, Cold Spring Harbor Symp. Quant. Biol. 30, 483 (1965).
    [Crossref]
  25. J. E. Dowling, Science 155, 273 (1967).
    [Crossref] [PubMed]
  26. L. Cervetto, Arch. Ital. Biol. 106, 194 (1968).
    [PubMed]
  27. R. Granit, Receptors and Sensory Perception (Yale University Press, New Haven, Conn., 1955), Ch. 5, p. 160.
  28. P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
    [Crossref]
  29. R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
    [Crossref] [PubMed]
  30. G. W. Hughes and L. Maffei, J. Neurophysiol. 29, 333 (1966).
    [PubMed]
  31. C. Enroth-Cugell and J. G. Robson, J. Physiol. (London) 187, 517 (1966).
  32. M. Clynes, Ann. N. Y. Acad. Sci. 92, 946 (1961).
    [Crossref] [PubMed]
  33. M. Clynes, Ann. N. Y. Acad. Sci. 98, 806 (1962). Also see Ann. N. Y. Acad. Sci. 156, Art. 2 (1969).
    [Crossref] [PubMed]

1968 (4)

D. Kahneman, Psychol. Bull. 70, 404 (1968).
[Crossref] [PubMed]

G. Sperling and M. M. Sondhi, J. Opt. Soc. Am. 58, 1133 (1968).
[Crossref] [PubMed]

R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
[Crossref] [PubMed]

L. Cervetto, Arch. Ital. Biol. 106, 194 (1968).
[PubMed]

1967 (2)

J. E. Dowling, Science 155, 273 (1967).
[Crossref] [PubMed]

M. Alpern and W. A. H. Rushton, J. Physiol. (London) 189, 519 (1967).

1966 (5)

G. Westheimer, Vision Res. 6, 669 (1966).
[Crossref] [PubMed]

B. Cleland and C. Enroth-Cugell, Acta Physiol. Scand. 68, 365 (1966).
[Crossref]

W. Rackensperger and O.-J. Grüsser, Experientia 22, 192 (1966).
[Crossref] [PubMed]

G. W. Hughes and L. Maffei, J. Neurophysiol. 29, 333 (1966).
[PubMed]

C. Enroth-Cugell and J. G. Robson, J. Physiol. (London) 187, 517 (1966).

1965 (4)

W. A. H. Rushton, J. Physiol. (London) 181, 645 (1965).

R. A. Cone, Cold Spring Harbor Symp. Quant. Biol. 30, 483 (1965).
[Crossref]

G. Sperling, J. Opt. Soc. Am. 55, 541 (1965).
[Crossref]

L. H. van der Tweel and H. F. E. Verduyn Lunel, EEG Clin. Neurophysiol. 18, 587 (1965).
[Crossref]

1964 (7)

M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
[Crossref] [PubMed]

G. Sperling, Doc. Ophthalmol. 18, 3 (1964).
[Crossref]

P. Gouras and R. D. Gunkel, Doc. Ophthalmol. 18, 137 (1964).
[Crossref]

D. H. Kelly, Doc. Ophthahlol. 18, 16 (1964).
[Crossref]

L. Matin, Am: J. Psychol. 77, 650 (1964).
[Crossref]

M. G. F. Fuortes and A. L. Hodgkin, J. Physiol. (London) 172, 239 (1964).

P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
[Crossref]

1963 (1)

G. Sperling, J. Opt. Soc. Am. 53, 520 (1963).

1962 (1)

M. Clynes, Ann. N. Y. Acad. Sci. 98, 806 (1962). Also see Ann. N. Y. Acad. Sci. 156, Art. 2 (1969).
[Crossref] [PubMed]

1961 (4)

M. Clynes, Ann. N. Y. Acad. Sci. 92, 946 (1961).
[Crossref] [PubMed]

D. H. Kelly, J. Opt. Soc. Am. 51, 422 (1961).
[Crossref] [PubMed]

L. H. van der Tweel, Ann. N. Y. Acad. Sci. 89, 829 (1961).
[Crossref] [PubMed]

R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
[Crossref]

1958 (1)

H. deLange, J. Opt. Soc. Am. 48, 777 (1958).
[Crossref]

1957 (1)

R. M. Boynton and G. L. Kandel, J. Opt. Soc. Am. 47, 275 (1957).
[Crossref] [PubMed]

1954 (1)

H. deLange, J. Opt. Soc. Am. 44, 380 (1954).
[Crossref]

Alpern, M.

M. Alpern and W. A. H. Rushton, J. Physiol. (London) 189, 519 (1967).

Boynton, R. M.

R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
[Crossref]

R. M. Boynton and G. L. Kandel, J. Opt. Soc. Am. 47, 275 (1957).
[Crossref] [PubMed]

Cervetto, L.

L. Cervetto, Arch. Ital. Biol. 106, 194 (1968).
[PubMed]

Cleland, B.

B. Cleland and C. Enroth-Cugell, Acta Physiol. Scand. 68, 365 (1966).
[Crossref]

Clynes, M.

M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
[Crossref] [PubMed]

M. Clynes, Ann. N. Y. Acad. Sci. 98, 806 (1962). Also see Ann. N. Y. Acad. Sci. 156, Art. 2 (1969).
[Crossref] [PubMed]

M. Clynes, Ann. N. Y. Acad. Sci. 92, 946 (1961).
[Crossref] [PubMed]

Cone, R. A.

R. A. Cone, Cold Spring Harbor Symp. Quant. Biol. 30, 483 (1965).
[Crossref]

deLange, H.

H. deLange, J. Opt. Soc. Am. 48, 777 (1958).
[Crossref]

H. deLange, J. Opt. Soc. Am. 44, 380 (1954).
[Crossref]

deVoe, R. G.

R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
[Crossref] [PubMed]

Dowling, J. E.

J. E. Dowling, Science 155, 273 (1967).
[Crossref] [PubMed]

Enroth-Cugell, C.

B. Cleland and C. Enroth-Cugell, Acta Physiol. Scand. 68, 365 (1966).
[Crossref]

C. Enroth-Cugell and J. G. Robson, J. Physiol. (London) 187, 517 (1966).

Fuortes, M. G. F.

M. G. F. Fuortes and A. L. Hodgkin, J. Physiol. (London) 172, 239 (1964).

Gouras, P.

P. Gouras and R. D. Gunkel, Doc. Ophthalmol. 18, 137 (1964).
[Crossref]

Granit, R.

R. Granit, Sensory Mechanisms of the Retina (Hafner Publishing Co., N. Y., 1963), Ch. 8, p. 145.

R. Granit, Receptors and Sensory Perception (Yale University Press, New Haven, Conn., 1955), Ch. 5, p. 160.

Grüsser, O.-J.

W. Rackensperger and O.-J. Grüsser, Experientia 22, 192 (1966).
[Crossref] [PubMed]

Gunkel, R. D.

P. Gouras and R. D. Gunkel, Doc. Ophthalmol. 18, 137 (1964).
[Crossref]

Hodgkin, A. L.

M. G. F. Fuortes and A. L. Hodgkin, J. Physiol. (London) 172, 239 (1964).

Hughes, G. W.

G. W. Hughes and L. Maffei, J. Neurophysiol. 29, 333 (1966).
[PubMed]

Ikeda, M.

R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
[Crossref]

Kahneman, D.

D. Kahneman, Psychol. Bull. 70, 404 (1968).
[Crossref] [PubMed]

Kandel, G. L.

R. M. Boynton and G. L. Kandel, J. Opt. Soc. Am. 47, 275 (1957).
[Crossref] [PubMed]

Kelly, D. H.

D. H. Kelly, Doc. Ophthahlol. 18, 16 (1964).
[Crossref]

D. H. Kelly, J. Opt. Soc. Am. 51, 422 (1961).
[Crossref] [PubMed]

Kohn, M.

M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
[Crossref] [PubMed]

Lifshitz, K.

M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
[Crossref] [PubMed]

Lunel, H. F. E. Verduyn

L. H. van der Tweel and H. F. E. Verduyn Lunel, EEG Clin. Neurophysiol. 18, 587 (1965).
[Crossref]

P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
[Crossref]

Maffei, L.

G. W. Hughes and L. Maffei, J. Neurophysiol. 29, 333 (1966).
[PubMed]

Matin, L.

L. Matin, Am: J. Psychol. 77, 650 (1964).
[Crossref]

Rackensperger, W.

W. Rackensperger and O.-J. Grüsser, Experientia 22, 192 (1966).
[Crossref] [PubMed]

Ripps, H.

R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
[Crossref] [PubMed]

Robson, J. G.

C. Enroth-Cugell and J. G. Robson, J. Physiol. (London) 187, 517 (1966).

Rushton, W. A. H.

M. Alpern and W. A. H. Rushton, J. Physiol. (London) 189, 519 (1967).

W. A. H. Rushton, J. Physiol. (London) 181, 645 (1965).

Sondhi, M. M.

G. Sperling and M. M. Sondhi, J. Opt. Soc. Am. 58, 1133 (1968).
[Crossref] [PubMed]

Sperling, G.

G. Sperling and M. M. Sondhi, J. Opt. Soc. Am. 58, 1133 (1968).
[Crossref] [PubMed]

G. Sperling, J. Opt. Soc. Am. 55, 541 (1965).
[Crossref]

G. Sperling, Doc. Ophthalmol. 18, 3 (1964).
[Crossref]

G. Sperling, J. Opt. Soc. Am. 53, 520 (1963).

Stoutenbeek, P.

P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
[Crossref]

Sturr, J. F.

R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
[Crossref]

van der Tweel, L. H.

L. H. van der Tweel and H. F. E. Verduyn Lunel, EEG Clin. Neurophysiol. 18, 587 (1965).
[Crossref]

P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
[Crossref]

L. H. van der Tweel, Ann. N. Y. Acad. Sci. 89, 829 (1961).
[Crossref] [PubMed]

Vaughan, Jr., H. G.

R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
[Crossref] [PubMed]

Westheimer, G.

G. Westheimer, Vision Res. 6, 669 (1966).
[Crossref] [PubMed]

Other (33)

G. Sperling, Doc. Ophthalmol. 18, 3 (1964).
[Crossref]

P. Gouras and R. D. Gunkel, Doc. Ophthalmol. 18, 137 (1964).
[Crossref]

B. Cleland and C. Enroth-Cugell, Acta Physiol. Scand. 68, 365 (1966).
[Crossref]

W. Rackensperger and O.-J. Grüsser, Experientia 22, 192 (1966).
[Crossref] [PubMed]

L. H. van der Tweel, Ann. N. Y. Acad. Sci. 89, 829 (1961).
[Crossref] [PubMed]

L. H. van der Tweel and H. F. E. Verduyn Lunel, EEG Clin. Neurophysiol. 18, 587 (1965).
[Crossref]

M. Clynes, M. Kohn, and K. Lifshitz, Ann. N. Y. Acad. Sci. 112, 468 (1964).
[Crossref] [PubMed]

D. Kahneman, Psychol. Bull. 70, 404 (1968).
[Crossref] [PubMed]

G. Sperling, J. Opt. Soc. Am. 55, 541 (1965).
[Crossref]

R. M. Boynton, J. F. Sturr, and M. Ikeda, J. Opt. Soc. Am. 51, 196 (1961).
[Crossref]

M. Alpern and W. A. H. Rushton, J. Physiol. (London) 189, 519 (1967).

H. deLange, J. Opt. Soc. Am. 44, 380 (1954).
[Crossref]

H. deLange, J. Opt. Soc. Am. 48, 777 (1958).
[Crossref]

G. Sperling and M. M. Sondhi, J. Opt. Soc. Am. 58, 1133 (1968).
[Crossref] [PubMed]

G. Sperling, J. Opt. Soc. Am. 53, 520 (1963).

D. H. Kelly, J. Opt. Soc. Am. 51, 422 (1961).
[Crossref] [PubMed]

D. H. Kelly, Doc. Ophthahlol. 18, 16 (1964).
[Crossref]

L. Matin, Am: J. Psychol. 77, 650 (1964).
[Crossref]

G. Westheimer, Vision Res. 6, 669 (1966).
[Crossref] [PubMed]

R. M. Boynton and G. L. Kandel, J. Opt. Soc. Am. 47, 275 (1957).
[Crossref] [PubMed]

R. Granit, Sensory Mechanisms of the Retina (Hafner Publishing Co., N. Y., 1963), Ch. 8, p. 145.

M. G. F. Fuortes and A. L. Hodgkin, J. Physiol. (London) 172, 239 (1964).

W. A. H. Rushton, J. Physiol. (London) 181, 645 (1965).

R. A. Cone, Cold Spring Harbor Symp. Quant. Biol. 30, 483 (1965).
[Crossref]

J. E. Dowling, Science 155, 273 (1967).
[Crossref] [PubMed]

L. Cervetto, Arch. Ital. Biol. 106, 194 (1968).
[PubMed]

R. Granit, Receptors and Sensory Perception (Yale University Press, New Haven, Conn., 1955), Ch. 5, p. 160.

P. Stoutenbeek, H. F. E. Verduyn Lunel, and L. H. van der Tweel, Doc. Ophthalmol. 18, 508 (1964).
[Crossref]

R. G. deVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968).
[Crossref] [PubMed]

G. W. Hughes and L. Maffei, J. Neurophysiol. 29, 333 (1966).
[PubMed]

C. Enroth-Cugell and J. G. Robson, J. Physiol. (London) 187, 517 (1966).

M. Clynes, Ann. N. Y. Acad. Sci. 92, 946 (1961).
[Crossref] [PubMed]

M. Clynes, Ann. N. Y. Acad. Sci. 98, 806 (1962). Also see Ann. N. Y. Acad. Sci. 156, Art. 2 (1969).
[Crossref] [PubMed]

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Figures (10)

Fig. 1
Fig. 1

Optical layout and electrical block diagram of apparatus. Meaning of symbols is explained in the text.

Fig. 2
Fig. 2

Test threshold as a function of the phase angle of the background cycle at the moment of the test flash. Observer MH. Frequency in hertz is shown on each graph. Modulation ratio m>99%, peak retinal illuminance, 1280 td. Points represent individual threshold settings. The sinusoid at the upper left illustrates the relative amplitudes of test and background stimuli.

Fig. 3
Fig. 3

Upper graph, maximum threshold energy as a function of frequency; lower graph, phase of background at which threshold was highest for each frequency. Points are means of 2–4 threshold settings. Modulation ratio m>99%, peak retinal illuminance, 1280 td. Observers DB (circles), MH (squares), AS (triangles). Values at 0 Hz are mean thresholds (n=12) measured with a 1280-td steady background. The arrow indicates the energy of a 2-msec test flash equal in amplitude to this background.

Fig. 4
Fig. 4

Initial maxima of threshold (means, n≥2) plotted relative to minima on a logarithmic ordinate, as functions of background frequency and modulation. All minima, if plotted, would fall on the zero line. Upper curves, m>99%; lower solid curves, m=50%; lower dashed curves, m=25%. Peak-to-peak change of retinal illuminance, 1280 td for all values of m. Observers DB (circles), MH (squares), AS (triangles).

Fig. 5
Fig. 5

Effect of steady light added to the 1280-td fluctuation. Test thresholds of observer DB are plotted as a function of the background phase at the moment of the test flash. Modulation percentages are indicated for each half of the figure, and frequency in hertz is shown in each graph. Bars indicate ranges between mean thresholds (n=6) measured with steady backgrounds equal to the extremes of the sinusoidal variation for each value of m. They are duplicated for comparison with dynamic thresholds obtained at the various frequencies.

Fig. 6
Fig. 6

Effect of steady light added to the 1280-td peak-to-peak background fluctuation. Data of observer MH. For explanation, see caption of Fig. 5.

Fig. 7
Fig. 7

Thresholds and threshold differences as functions of background frequency: (a) initial maxima, (b) initial maxima minus minima. The minima (c) are the averages of settings obtained at or near the end of the first run and the beginning of the second run in each experimental session. Upper panels, observer DB; lower panels, MH. Parameter is modulation ratio: m>99% (circles), 50% (squares), 25% (triangles). Peak-to-peak change of retinal illuminance, 1280 td.

Fig. 8
Fig. 8

Thresholds and threshold differences as functions of background frequency: (a) late maxima, (b) late maxima minus the minima plotted in Fig. 7(c). Upper panels, observer DB; lower panels, MH. Parameter is modulation ratio, m=50% (squares), 25% (triangles). Peak-to-peak change of retinal illuminance, 1280 td.

Fig. 9
Fig. 9

Threshold functions of observer AS obtained with a low-level background. Frequency in hertz is shown in each graph. Modulation ratio m>99%, peak retinal illuminance 128 td. The bar in each panel indicates the range between mean thresholds (n=8) measured with steady backgrounds equal to the extremes of the sinusoidal variation. Points represent individual threshold settings.

Fig. 10
Fig. 10

Threshold maxima (upper curves) and minima of observer AS as functions of background frequency. Modulation ratio m>99%, peak retinal illuminance 128 td [part (a)], 1280 td [part (b)]. Each point is the mean of two threshold settings.

Tables (2)

Tables Icon

Table I Maximal and minimal threshold values, their differences and their ratios, for m>99% (peak-to-peak amplitude, 1280 td).

Tables Icon

Table II Phase angles of sinusoid coincident with maxima and minima of threshold curves.