Abstract

It has been suggested that local variations of retinal sensitivity may be responsible for elevating the threshold in pattern-adaptation experiments of the Blakemore–Campbell type. Subjects are unable to scan high-contrast gratings uniformly enough to eliminate this possibility. To control this effect, we performed grating-adaptation experiments under stabilized-image conditions, while both adapting and test targets were moved at retinal velocities determined by the experimenter. By means of an afterimage technique, we also measured the strength of the retinal sensitivity mask that forms under these conditions. Varying the spatial frequency and velocity of the adapting stimulus, we inferred the spatial and temporal properties of the principal mechanism that contributes to the afterimage. We found that the Blakemore–Campbell effect persists at adapting velocities that are fast enough to rule out local variations of retinal sensitivity. More surprisingly, even the clearly visible afterimages that occur at a retinal velocity of 0.1 deg/s seem to have no effect on pattern adaptation. (Sensitivity masking can raise the adapted threshold, but only at adapting velocities slower than normal eye movements.) By manipulating the image velocity, we were able to shift the spatial frequencies of some threshold-elevation curves, but these shifts were not great enough to suggest that velocity tuning plays important role in pattern adaptation.

© 1980 Optical Society of America

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References

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  1. D. H. Kelly, “Motion and vision. I. Stabilized images of stationary gratings,” J. Opt. Soc. Am. 69, 1266–1274 (1979).
    [Crossref] [PubMed]
  2. D. H. Kelly, “Visual contrast sensitivity,” Opt. Acta 24, 107–129 (1977).
    [Crossref]
  3. D. H. Kelly, “Motion and vision. II. Stabilized spatio-temporal threshold surface,” J. Opt. Soc. Am. 69, 1340–1349 (1979).
    [Crossref] [PubMed]
  4. C. Blakemore and F. W. Campbell, “On the existence of neurones in the human visual system selectively sensitive to the orientation and size of retinal images,” J. Physiol. (London) 203, 237–260 (1969).
  5. R. M. Jones and U. Tulunay-Keesey, “Local retinal adaptation and spatial frequency channels,” Vision Res. 15, 1239–1244 (1975).
    [Crossref] [PubMed]
  6. R. M. Jones and U. Tulunay-Keesey, “Phase selectivity of spatial frequency channels,” J. Opt. Soc. Am. 70, 66–70 (1980).
    [Crossref] [PubMed]
  7. N. Graham, “Spatial frequency channels in the human visual system: Effects of luminance and pattern drift rate,” Vision Res. 12, 53–68 (1972).
    [Crossref] [PubMed]
  8. D. J. Tolhurst, “Separate channels for the analysis of the shape and the movements of a moving visual stimulus,” J. Physiol. (London) 231, 385–402 (1973).
  9. L. E. Arend and A. A. Skavenski, “Free scanning of gratings produces patterned retinal exposure,” Vision Res. 19, 1413–1419 (1979).
    [Crossref] [PubMed]
  10. Conversely, if the duration of a transient stimulus is short enough to “freeze” any natural eye movement, then stabilizing the retinal image obviously can have no effect. In the intermediate range of longer transients, the effect of stabilization seems to be variable and difficult to interpret. By comparison, steady-state stimulation is a much more revealing technique for exploring stabilized-image effects.
  11. D. H. Kelly and R. E. Savoie, “A study of sine-wave contrast sensitivity by two psychophysical methods,” Percept. Psychophys. 14, 313–318 (1973).
    [Crossref]
  12. Samples of our eye-movement data were originally included in this paper, but were subsequently deleted at the suggestion of a referee.
  13. D. H. Fender (personal communication).
  14. J. J. Koenderink, “Contrast enhancement and the negative afterimage,” J. Opt. Soc. Am. 62, 685–689 (1972).
    [Crossref] [PubMed]
  15. D. H. Kelly, “Theory of flicker and transient responses. I. Uniform fields,” J. Opt. Soc. Am. 61, 537–546 (1971).
    [Crossref] [PubMed]
  16. D. H. Kelly, “Adaptation effects on spatio-temporal sine-wave thresholds,” Vision Res. 12, 89–101 (1972).
    [Crossref] [PubMed]
  17. The bandwidths shown in Fig. 5 may appear broader than those of Refs. 4, 5, and others, but this is due to a different way of plotting the data. Blakemore and Campbell subtracted 1 from their threshold elevation ratios and plotted the result on a logarithmic scale, whereas we plot the log of the ratio directly.
  18. This effect has been studied in detail by Karen DeValois, “Spatial frequency adaptation can enhance contrast sensitivity,” Vision Res. 17, 1057–1065 (1977).
    [Crossref]
  19. See Ref. 3, Fig. 11.
  20. F. L. Van Nes, J. J. Koenderink, H. Nas, and M. A. Bouman, “Spatiotemporal modulation transfer in the human eye,” J. Opt. Soc. Am. 57, 1082–1088 (1967).
    [Crossref] [PubMed]
  21. D. H. Kelly, “Frequency doubling in visual responses,” J. Opt. Soc. Am. 56, 1628–1633 (1966).
    [Crossref]
  22. V. Virsu and P. Laurinen, “Long-lasting afterimages caused by neural adaptation,” Vision Res. 17, 853–860 (1977).
    [Crossref] [PubMed]

1980 (1)

1979 (3)

1977 (3)

This effect has been studied in detail by Karen DeValois, “Spatial frequency adaptation can enhance contrast sensitivity,” Vision Res. 17, 1057–1065 (1977).
[Crossref]

V. Virsu and P. Laurinen, “Long-lasting afterimages caused by neural adaptation,” Vision Res. 17, 853–860 (1977).
[Crossref] [PubMed]

D. H. Kelly, “Visual contrast sensitivity,” Opt. Acta 24, 107–129 (1977).
[Crossref]

1975 (1)

R. M. Jones and U. Tulunay-Keesey, “Local retinal adaptation and spatial frequency channels,” Vision Res. 15, 1239–1244 (1975).
[Crossref] [PubMed]

1973 (2)

D. J. Tolhurst, “Separate channels for the analysis of the shape and the movements of a moving visual stimulus,” J. Physiol. (London) 231, 385–402 (1973).

D. H. Kelly and R. E. Savoie, “A study of sine-wave contrast sensitivity by two psychophysical methods,” Percept. Psychophys. 14, 313–318 (1973).
[Crossref]

1972 (3)

J. J. Koenderink, “Contrast enhancement and the negative afterimage,” J. Opt. Soc. Am. 62, 685–689 (1972).
[Crossref] [PubMed]

D. H. Kelly, “Adaptation effects on spatio-temporal sine-wave thresholds,” Vision Res. 12, 89–101 (1972).
[Crossref] [PubMed]

N. Graham, “Spatial frequency channels in the human visual system: Effects of luminance and pattern drift rate,” Vision Res. 12, 53–68 (1972).
[Crossref] [PubMed]

1971 (1)

1969 (1)

C. Blakemore and F. W. Campbell, “On the existence of neurones in the human visual system selectively sensitive to the orientation and size of retinal images,” J. Physiol. (London) 203, 237–260 (1969).

1967 (1)

1966 (1)

Arend, L. E.

L. E. Arend and A. A. Skavenski, “Free scanning of gratings produces patterned retinal exposure,” Vision Res. 19, 1413–1419 (1979).
[Crossref] [PubMed]

Blakemore, C.

C. Blakemore and F. W. Campbell, “On the existence of neurones in the human visual system selectively sensitive to the orientation and size of retinal images,” J. Physiol. (London) 203, 237–260 (1969).

Bouman, M. A.

Campbell, F. W.

C. Blakemore and F. W. Campbell, “On the existence of neurones in the human visual system selectively sensitive to the orientation and size of retinal images,” J. Physiol. (London) 203, 237–260 (1969).

DeValois, Karen

This effect has been studied in detail by Karen DeValois, “Spatial frequency adaptation can enhance contrast sensitivity,” Vision Res. 17, 1057–1065 (1977).
[Crossref]

Fender, D. H.

D. H. Fender (personal communication).

Graham, N.

N. Graham, “Spatial frequency channels in the human visual system: Effects of luminance and pattern drift rate,” Vision Res. 12, 53–68 (1972).
[Crossref] [PubMed]

Jones, R. M.

R. M. Jones and U. Tulunay-Keesey, “Phase selectivity of spatial frequency channels,” J. Opt. Soc. Am. 70, 66–70 (1980).
[Crossref] [PubMed]

R. M. Jones and U. Tulunay-Keesey, “Local retinal adaptation and spatial frequency channels,” Vision Res. 15, 1239–1244 (1975).
[Crossref] [PubMed]

Kelly, D. H.

Koenderink, J. J.

Laurinen, P.

V. Virsu and P. Laurinen, “Long-lasting afterimages caused by neural adaptation,” Vision Res. 17, 853–860 (1977).
[Crossref] [PubMed]

Nas, H.

Savoie, R. E.

D. H. Kelly and R. E. Savoie, “A study of sine-wave contrast sensitivity by two psychophysical methods,” Percept. Psychophys. 14, 313–318 (1973).
[Crossref]

Skavenski, A. A.

L. E. Arend and A. A. Skavenski, “Free scanning of gratings produces patterned retinal exposure,” Vision Res. 19, 1413–1419 (1979).
[Crossref] [PubMed]

Tolhurst, D. J.

D. J. Tolhurst, “Separate channels for the analysis of the shape and the movements of a moving visual stimulus,” J. Physiol. (London) 231, 385–402 (1973).

Tulunay-Keesey, U.

R. M. Jones and U. Tulunay-Keesey, “Phase selectivity of spatial frequency channels,” J. Opt. Soc. Am. 70, 66–70 (1980).
[Crossref] [PubMed]

R. M. Jones and U. Tulunay-Keesey, “Local retinal adaptation and spatial frequency channels,” Vision Res. 15, 1239–1244 (1975).
[Crossref] [PubMed]

Van Nes, F. L.

Virsu, V.

V. Virsu and P. Laurinen, “Long-lasting afterimages caused by neural adaptation,” Vision Res. 17, 853–860 (1977).
[Crossref] [PubMed]

J. Opt. Soc. Am. (7)

J. Physiol. (London) (2)

D. J. Tolhurst, “Separate channels for the analysis of the shape and the movements of a moving visual stimulus,” J. Physiol. (London) 231, 385–402 (1973).

C. Blakemore and F. W. Campbell, “On the existence of neurones in the human visual system selectively sensitive to the orientation and size of retinal images,” J. Physiol. (London) 203, 237–260 (1969).

Opt. Acta (1)

D. H. Kelly, “Visual contrast sensitivity,” Opt. Acta 24, 107–129 (1977).
[Crossref]

Percept. Psychophys. (1)

D. H. Kelly and R. E. Savoie, “A study of sine-wave contrast sensitivity by two psychophysical methods,” Percept. Psychophys. 14, 313–318 (1973).
[Crossref]

Vision Res. (6)

This effect has been studied in detail by Karen DeValois, “Spatial frequency adaptation can enhance contrast sensitivity,” Vision Res. 17, 1057–1065 (1977).
[Crossref]

D. H. Kelly, “Adaptation effects on spatio-temporal sine-wave thresholds,” Vision Res. 12, 89–101 (1972).
[Crossref] [PubMed]

R. M. Jones and U. Tulunay-Keesey, “Local retinal adaptation and spatial frequency channels,” Vision Res. 15, 1239–1244 (1975).
[Crossref] [PubMed]

L. E. Arend and A. A. Skavenski, “Free scanning of gratings produces patterned retinal exposure,” Vision Res. 19, 1413–1419 (1979).
[Crossref] [PubMed]

N. Graham, “Spatial frequency channels in the human visual system: Effects of luminance and pattern drift rate,” Vision Res. 12, 53–68 (1972).
[Crossref] [PubMed]

V. Virsu and P. Laurinen, “Long-lasting afterimages caused by neural adaptation,” Vision Res. 17, 853–860 (1977).
[Crossref] [PubMed]

Other (5)

Conversely, if the duration of a transient stimulus is short enough to “freeze” any natural eye movement, then stabilizing the retinal image obviously can have no effect. In the intermediate range of longer transients, the effect of stabilization seems to be variable and difficult to interpret. By comparison, steady-state stimulation is a much more revealing technique for exploring stabilized-image effects.

The bandwidths shown in Fig. 5 may appear broader than those of Refs. 4, 5, and others, but this is due to a different way of plotting the data. Blakemore and Campbell subtracted 1 from their threshold elevation ratios and plotted the result on a logarithmic scale, whereas we plot the log of the ratio directly.

See Ref. 3, Fig. 11.

Samples of our eye-movement data were originally included in this paper, but were subsequently deleted at the suggestion of a referee.

D. H. Fender (personal communication).

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Figures (7)

FIG. 1
FIG. 1

Schematic diagram of the spatiotemporal frequency domain. In log–log coordinates, the locus of any constant-velocity stimulus is a straight line oriented at 45 deg. Three examples are shown, for 0.1, 3.2, and 100 deg/s. The two lower velocities represent stimuli used in Secs. IV and V. The shaded area shows the region where afterimages are readily detectable with stabilized viewing. (A retinal image velocity of 0.1 deg/s yields essentially the same contrast sensitivity function as that obtained with natural eye movements.)

FIG. 2
FIG. 2

Threshold-afterimage measurements of sensitivity masks formed by stabilized gratings. For the upper solid curve (open squares), the gratings were stationary; for the lower solid curve (open circles), they were moving at a constant velocity of 0.15 deg/s. Dashed curves show theoretical extrapolations described in the text.

FIG. 3
FIG. 3

Ratio of the two sensitivity masks shown in Fig. 2. Dashed line indicates a slope of 45° (in log–log coordinates).

FIG. 4
FIG. 4

Upper curves show the stabilized contrast sensitivity with (open circles) and without (filled circles) adaptation to a high-contrast, 6-c/deg grating. The adapting stimulus and all test stimuli were moving at 0.15 deg/s. The lower curve shows the ratio of the thresholds obtained under these two conditions.

FIG. 5
FIG. 5

Stabilized threshold–elevation curves showing the effect of changing the spatial frequency of the adapting grating. Adapting frequencies were 1 (triangles), 3 (squares), and 6 c/deg (circles). Other conditions, same as in Fig. 4.

FIG. 6
FIG. 6

Effect of changing the velocity of a 1-c/deg adapting grating; test velocity, 0.1 deg/s. Adapting velocities from 0.1 (open squares) to 3.2 deg/s (filled circles) give the same threshold elevation curve. However, threshold elevation is greatly increased when the adapting grating is stationary (open circles).

FIG. 7
FIG. 7

Effect of adapting and test velocities of 0.1 and 3.2 deg/s on threshold elevation for a 3-c/deg adapting grating. The peak is shifted toward lower frequencies when the adapting velocity is higher than the test velocity and toward higher frequencies when it is lower.