Abstract

We found that inspecting a sine-wave grating elevated threshold for spatial-frequency discrimination as it does for contrast detection, but discrimination threshold was maximally elevated at about twice the adapting frequency, where detection threshold was little affected; and detection threshold was maximally elevated at the adapting frequency, where discrimination threshold was not elevated at all. Orientation tuning was roughly similar for contrast and for discrimination threshold elevations; elevations fell by half at between 7 and 17 deg from the adapting orientation. We compared our findings with the predictions of three models of discrimination: (1) The data are inconsistent with the idea that the most strongly stimulated channels are the most important channels for discrimination. (2) With an additional assumption, the Hirsch—Hylton scaled-lattice model could account for our finding that discrimination threshold elevations are asymmetric. (3) With no additional assumptions, the idea that discriminati n is determined by the relative activities of multiple overlapping spatial-frequency channels or sizetuned neurons can account for our finding that discrimination thresholds are asymmetric. We propose a physiologically based discrimination model: Asymmetrically tuned cortical cells feed a ratio-tuned neural mechanism whose properties are formally analogous to those of ratio-tuned neurons that have recently been found in cat visual cortex. The linear relation between firing frequency and contrast can explain why discrimination threshold is substantially independent of contrast.

© 1983 Optical Society of America

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  1. F. W. Campbell and J. G. Robson, "Applications of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).
  2. C. B. Blakemore and F. W. Campbell, "On the existence of neurons in the human visual system selectively sensitive to the orientation and size of retinal image," J. Physiol. 203, 237–260 (1969).
  3. N. Graham, "Spatial frequency channels in human vision: detecting edges without edge detectors," in Visual Coding and Adaptability, C. S. Harris, ed. (Erlbaum, Hillsdale, N.J., 1980).
  4. O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.
  5. D. H. Kelly and C. A. Burbeck, "Critical problems in spatial vision," in Critical Reuiews in Bioengineering (CRC, Boca Raton, Fla., 1983) (to be published).
  6. F. W. Campbell, J. Nachmias, and J. Jukes, "Spatial-frequency discrimination in human vision," J. Opt. Soc. Am. 60, 555–559 (1970).
  7. J. Hirsch and R. Hylton, "Limits of spatial frequency discrimination as evidence of neural interpolation," J. Opt. Soc. Am. 72, 1367–1374 (1982).
  8. A. B. Watson and J. G. Robson, "Discrimination at threshold: labelled detectors in human vision," Vision Res. 21, 1115–1122 (1981).
  9. H. R. Wilson and D. J. Gelb, "A modified line element theory for spatial frequency and width discrimination," J. Opt. Soc. Am. (to be published).
  10. D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).
  11. D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).
  12. I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).
  13. R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).
  14. C. Blakemore and J. Nachmias, "The orientational specificity of two visual aftereffects," J. Physiol. 213, 157–174 (1971).
  15. J. A. Movshon and C. Blakemore, "Orientation specificity and spatial selectivity in human vision," Perception 2, 53–60 (1973).
  16. F. W. Campbell and J. J. Kulikowski, "Orientation selectivity of the human visual system," J. Physiol. 187, 437–445 (1966).
  17. Caveats include the following: (1) 1(o take the bandwidth of the adaptation effect as the excitatory bandwidth of a channel implies that selective adaptation is a desensitization that is due to prolonged excitation, and there is some evidence that inhibition is involved.4 (2) In any case, the angular bandwidth of the threshold elevation is not necessarily the same as the angular bandwidth of a channel, since different channels may determine threshold before and after adaptation.5
  18. This way of conceptually linking detection and discrimination is not restricted to spatial vision and has been applied to several visual modalities.19 In the context of color vision, Boynton20 discusses how the approach can be formulated either as a line element model, in which central neural processing has direct access to channel activities, or as an opponent process model, in which opponent mechanisms intervene between channels and subsequent processing. If they are both linear, the two formulations are equivalent. Historically, these two formulations of the relative activity approach have been influential and quite successful in color-vision research.20 More recently, this same concept has been invoked to account for acute discrimination between different orientations21 and acute discrimination of motion in depth.22
  19. D. Regan, "Visual information channeling in normal and disordered vision," Psychol. Rev. 89, 407–444 (1982).
  20. R. M. Boynton, Human Color Vision (Holt, Rinehart & Winston, New York, 1979).
  21. G. Westheimer, K. Shimamura, and S. P. McKee, "Interference with line-orientation sensitivity," J. Opt. Soc. Am. 66, 332–338 (1976).
  22. K. I. Beverley and D. Regan, "The relation between discrimination and sensitivity in the perception of motion in depth," J. Physiol. 249, 387–398 (1975).
  23. H. Wilson, Department of Biophysics and Theoretical Biology, University of Chicago, Chicago, Ill. 60637 (personal communication).
  24. M. Cynader and D. Regan, "Neurons in cat parastriate cortex sensitive to the direction of motion in three-dimensional space," J. Physiol. 274, 549–569 (1978).
  25. D. Regan and M. Cynader, "Neurons in cat visual cortex tuned to the direction of motion in depth: effect of stimulus speed," Invest. Ophthalmol. Vis. Sci. 22, 535–550 (1982).
  26. R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.
  27. S1 and S2 would be the excitatory center/inhibitory surround type of neuron26 for pairs of dark lines and inhibitory center/excitatory surround26 for bright line pairs.
  28. G. F. Cooper and J. G. Robson, "Application of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).
  29. V. A. Movshon and D. J. Tolhurst, "On the response linearity of cells in the cat visual cortex," J. Physiol. 249, 56–57P (1975).
  30. A. F. Dean, "The relationship between response amplitude and contrast for cat striate cortical neurones," J. Physiol. 318, 413–427 (1981).
  31. J. Hirsch and R. Hylton, Department of Ophthalmology, Yale University, New Haven, Conn. 06520 (personal communication).
  32. D. Regan, "Spatial frequency mechanisms in human vision: electrophysiological evidence," Vision Res. (to be published).
  33. H. Wilson, Department of Biophysics and Theoretical Biology, University of Chicago, Chicago, Ill. 60637 (personal communication).
  34. H. R. Wilson, D. K. McFarlane, and G. C. Phillips, "Spatial tuning of orientation selective units estimated by oblique masking," Vision Res. (to be published).
  35. P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).
  36. J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).
  37. D. J. Tolhurst and I. D. Thompson, "On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat," Proc. R. Soc. Lond. Ser. B 213, 183–199 (1981).
  38. F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).
  39. Presumably the most important among the neurons contributing to discrimination are those with the steepest slopes. However, it is not clear from published data on the spatial-frequency tuning of cortical cells whether the reason that neither symmetrically tuned neurons nor cells with steeper high- than low-frequency slopes seem to contribute importantly to discrimination is merely that these kinds of tuning curves are not associated with the steepest slopes.

1983 (1)

D. H. Kelly and C. A. Burbeck, "Critical problems in spatial vision," in Critical Reuiews in Bioengineering (CRC, Boca Raton, Fla., 1983) (to be published).

1982 (4)

J. Hirsch and R. Hylton, "Limits of spatial frequency discrimination as evidence of neural interpolation," J. Opt. Soc. Am. 72, 1367–1374 (1982).

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

D. Regan, "Visual information channeling in normal and disordered vision," Psychol. Rev. 89, 407–444 (1982).

D. Regan and M. Cynader, "Neurons in cat visual cortex tuned to the direction of motion in depth: effect of stimulus speed," Invest. Ophthalmol. Vis. Sci. 22, 535–550 (1982).

1981 (3)

A. B. Watson and J. G. Robson, "Discrimination at threshold: labelled detectors in human vision," Vision Res. 21, 1115–1122 (1981).

A. F. Dean, "The relationship between response amplitude and contrast for cat striate cortical neurones," J. Physiol. 318, 413–427 (1981).

D. J. Tolhurst and I. D. Thompson, "On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat," Proc. R. Soc. Lond. Ser. B 213, 183–199 (1981).

1980 (1)

N. Graham, "Spatial frequency channels in human vision: detecting edges without edge detectors," in Visual Coding and Adaptability, C. S. Harris, ed. (Erlbaum, Hillsdale, N.J., 1980).

1979 (2)

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).

1978 (4)

J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).

M. Cynader and D. Regan, "Neurons in cat parastriate cortex sensitive to the direction of motion in three-dimensional space," J. Physiol. 274, 549–569 (1978).

R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.

O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.

1977 (1)

D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).

1976 (2)

P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).

G. Westheimer, K. Shimamura, and S. P. McKee, "Interference with line-orientation sensitivity," J. Opt. Soc. Am. 66, 332–338 (1976).

1975 (2)

V. A. Movshon and D. J. Tolhurst, "On the response linearity of cells in the cat visual cortex," J. Physiol. 249, 56–57P (1975).

K. I. Beverley and D. Regan, "The relation between discrimination and sensitivity in the perception of motion in depth," J. Physiol. 249, 387–398 (1975).

1973 (1)

J. A. Movshon and C. Blakemore, "Orientation specificity and spatial selectivity in human vision," Perception 2, 53–60 (1973).

1971 (1)

C. Blakemore and J. Nachmias, "The orientational specificity of two visual aftereffects," J. Physiol. 213, 157–174 (1971).

1970 (1)

1969 (2)

C. B. Blakemore and F. W. Campbell, "On the existence of neurons in the human visual system selectively sensitive to the orientation and size of retinal image," J. Physiol. 203, 237–260 (1969).

F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).

1968 (2)

F. W. Campbell and J. G. Robson, "Applications of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

G. F. Cooper and J. G. Robson, "Application of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

1966 (1)

F. W. Campbell and J. J. Kulikowski, "Orientation selectivity of the human visual system," J. Physiol. 187, 437–445 (1966).

Albrecht, D. G.

R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.

Atkinson, J.

O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.

Bartol, S.

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

Beverley, K. I.

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

K. I. Beverley and D. Regan, "The relation between discrimination and sensitivity in the perception of motion in depth," J. Physiol. 249, 387–398 (1975).

Blakemore, C.

J. A. Movshon and C. Blakemore, "Orientation specificity and spatial selectivity in human vision," Perception 2, 53–60 (1973).

C. Blakemore and J. Nachmias, "The orientational specificity of two visual aftereffects," J. Physiol. 213, 157–174 (1971).

Blakemore, C. B.

C. B. Blakemore and F. W. Campbell, "On the existence of neurons in the human visual system selectively sensitive to the orientation and size of retinal image," J. Physiol. 203, 237–260 (1969).

Bodis-Wollner, I.

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

Boynton, R. M.

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

Braddick, O.

O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.

Burbeck, C. A.

D. H. Kelly and C. A. Burbeck, "Critical problems in spatial vision," in Critical Reuiews in Bioengineering (CRC, Boca Raton, Fla., 1983) (to be published).

Campbell, F. W.

R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).

O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.

F. W. Campbell, J. Nachmias, and J. Jukes, "Spatial-frequency discrimination in human vision," J. Opt. Soc. Am. 60, 555–559 (1970).

F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).

C. B. Blakemore and F. W. Campbell, "On the existence of neurons in the human visual system selectively sensitive to the orientation and size of retinal image," J. Physiol. 203, 237–260 (1969).

F. W. Campbell and J. G. Robson, "Applications of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

F. W. Campbell and J. J. Kulikowski, "Orientation selectivity of the human visual system," J. Physiol. 187, 437–445 (1966).

Cooper, G. F.

F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).

G. F. Cooper and J. G. Robson, "Application of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

Cynader, M.

D. Regan and M. Cynader, "Neurons in cat visual cortex tuned to the direction of motion in depth: effect of stimulus speed," Invest. Ophthalmol. Vis. Sci. 22, 535–550 (1982).

M. Cynader and D. Regan, "Neurons in cat parastriate cortex sensitive to the direction of motion in three-dimensional space," J. Physiol. 274, 549–569 (1978).

De Valois, R. L.

R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.

Dean, A. F.

A. F. Dean, "The relationship between response amplitude and contrast for cat striate cortical neurones," J. Physiol. 318, 413–427 (1981).

Enroth-Cugell, C.

F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).

Finlay, B. L.

P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).

Gelb, D. J.

H. R. Wilson and D. J. Gelb, "A modified line element theory for spatial frequency and width discrimination," J. Opt. Soc. Am. (to be published).

Graham, N.

N. Graham, "Spatial frequency channels in human vision: detecting edges without edge detectors," in Visual Coding and Adaptability, C. S. Harris, ed. (Erlbaum, Hillsdale, N.J., 1980).

Hendley, C. D.

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

Hirsch, J.

J. Hirsch and R. Hylton, "Limits of spatial frequency discrimination as evidence of neural interpolation," J. Opt. Soc. Am. 72, 1367–1374 (1982).

J. Hirsch and R. Hylton, Department of Ophthalmology, Yale University, New Haven, Conn. 06520 (personal communication).

Hylton, R.

J. Hirsch and R. Hylton, "Limits of spatial frequency discrimination as evidence of neural interpolation," J. Opt. Soc. Am. 72, 1367–1374 (1982).

J. Hirsch and R. Hylton, Department of Ophthalmology, Yale University, New Haven, Conn. 06520 (personal communication).

Jukes, J.

Kelly, D. H.

D. H. Kelly and C. A. Burbeck, "Critical problems in spatial vision," in Critical Reuiews in Bioengineering (CRC, Boca Raton, Fla., 1983) (to be published).

Kulikowski, J. J.

F. W. Campbell and J. J. Kulikowski, "Orientation selectivity of the human visual system," J. Physiol. 187, 437–445 (1966).

McFarlane, D. K.

H. R. Wilson, D. K. McFarlane, and G. C. Phillips, "Spatial tuning of orientation selective units estimated by oblique masking," Vision Res. (to be published).

McKee, S. P.

Movshon, J. A.

J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).

J. A. Movshon and C. Blakemore, "Orientation specificity and spatial selectivity in human vision," Perception 2, 53–60 (1973).

Movshon, V. A.

V. A. Movshon and D. J. Tolhurst, "On the response linearity of cells in the cat visual cortex," J. Physiol. 249, 56–57P (1975).

Murray, T. J.

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).

Mylin, L. H.

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

Nachmias, J.

C. Blakemore and J. Nachmias, "The orientational specificity of two visual aftereffects," J. Physiol. 213, 157–174 (1971).

F. W. Campbell, J. Nachmias, and J. Jukes, "Spatial-frequency discrimination in human vision," J. Opt. Soc. Am. 60, 555–559 (1970).

Phillips, G. C.

H. R. Wilson, D. K. McFarlane, and G. C. Phillips, "Spatial tuning of orientation selective units estimated by oblique masking," Vision Res. (to be published).

Regan, D.

D. Regan and M. Cynader, "Neurons in cat visual cortex tuned to the direction of motion in depth: effect of stimulus speed," Invest. Ophthalmol. Vis. Sci. 22, 535–550 (1982).

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

D. Regan, "Visual information channeling in normal and disordered vision," Psychol. Rev. 89, 407–444 (1982).

M. Cynader and D. Regan, "Neurons in cat parastriate cortex sensitive to the direction of motion in three-dimensional space," J. Physiol. 274, 549–569 (1978).

D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).

K. I. Beverley and D. Regan, "The relation between discrimination and sensitivity in the perception of motion in depth," J. Physiol. 249, 387–398 (1975).

D. Regan, "Spatial frequency mechanisms in human vision: electrophysiological evidence," Vision Res. (to be published).

Robson, J. G.

A. B. Watson and J. G. Robson, "Discrimination at threshold: labelled detectors in human vision," Vision Res. 21, 1115–1122 (1981).

F. W. Campbell and J. G. Robson, "Applications of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

G. F. Cooper and J. G. Robson, "Application of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

Schiller, P. H.

P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).

Shimamura, K.

Silver, R.

D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).

Thompson, I. D.

D. J. Tolhurst and I. D. Thompson, "On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat," Proc. R. Soc. Lond. Ser. B 213, 183–199 (1981).

J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).

Thorell, L. G.

R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.

Thornton, J.

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

Tolhurst, D. J.

D. J. Tolhurst and I. D. Thompson, "On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat," Proc. R. Soc. Lond. Ser. B 213, 183–199 (1981).

J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).

V. A. Movshon and D. J. Tolhurst, "On the response linearity of cells in the cat visual cortex," J. Physiol. 249, 56–57P (1975).

Volman, S. F.

P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).

Watson, A. B.

A. B. Watson and J. G. Robson, "Discrimination at threshold: labelled detectors in human vision," Vision Res. 21, 1115–1122 (1981).

Westheimer, G.

Wilkinson, I. M. S.

R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).

Wilson, H.

H. Wilson, Department of Biophysics and Theoretical Biology, University of Chicago, Chicago, Ill. 60637 (personal communication).

Wilson, H. R.

H. R. Wilson and D. J. Gelb, "A modified line element theory for spatial frequency and width discrimination," J. Opt. Soc. Am. (to be published).

H. R. Wilson, D. K. McFarlane, and G. C. Phillips, "Spatial tuning of orientation selective units estimated by oblique masking," Vision Res. (to be published).

Zimmern, R. L.

R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).

Ann. Neurol. (1)

I. Bodis-Wollner, C. D. Hendley, L. H. Mylin, and J. Thornton, "Visual evoked potentials and the visuogram in multiple sclerosis," Ann. Neurol. 5, 40–47 (1979).

Brain (2)

D. Regan, S. Bartol, T. J. Murray, and K. I. Beverley, "Spatial frequency discrimination in normal vision and in patients with multiple sclerosis," Brain 104, 735–754 (1982).

D. Regan, R. Silver, and T. J. Murray, "Visual acuity and contrast sensitivity in multiple sclerosis: hidden visual loss," Brain 100, 563–579 (1977).

Invest. Ophthalmol. Vis. Sci. (1)

D. Regan and M. Cynader, "Neurons in cat visual cortex tuned to the direction of motion in depth: effect of stimulus speed," Invest. Ophthalmol. Vis. Sci. 22, 535–550 (1982).

J. Neurol. Neurosurg. Psychiat. (1)

R. L. Zimmern, F. W. Campbell, and I. M. S. Wilkinson, "Subtle disturbances of vision after optic neuritis elicited by studying contrast sensitivity," J. Neurol. Neurosurg. Psychiat. 42, 407–412 (1979).

J. Neurophysiol. (1)

P. H. Schiller, B. L. Finlay, and S. F. Volman, "Quantitative studies of single-cell properties in monkey striate cortex. III. Spatial frequency," J. Neurophysiol. 39, 1334–1351 (1976).

J. Opt. Soc. Am. (3)

J. Physiol. (11)

M. Cynader and D. Regan, "Neurons in cat parastriate cortex sensitive to the direction of motion in three-dimensional space," J. Physiol. 274, 549–569 (1978).

F. W. Campbell and J. G. Robson, "Applications of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

C. B. Blakemore and F. W. Campbell, "On the existence of neurons in the human visual system selectively sensitive to the orientation and size of retinal image," J. Physiol. 203, 237–260 (1969).

C. Blakemore and J. Nachmias, "The orientational specificity of two visual aftereffects," J. Physiol. 213, 157–174 (1971).

F. W. Campbell and J. J. Kulikowski, "Orientation selectivity of the human visual system," J. Physiol. 187, 437–445 (1966).

J. A. Movshon, I. D. Thompson, and D. J. Tolhurst, "Spatial and temporal contrast sensitivity of neurons in areas 17 and 18 of the cat's visual cortex," J. Physiol. 283, 101–120 (1978).

F. W. Campbell, G. F. Cooper, and C. Enroth-Cugell, "The spatial selectivity of the visual cells of the cat," J. Physiol. 203, 223–235 (1969).

K. I. Beverley and D. Regan, "The relation between discrimination and sensitivity in the perception of motion in depth," J. Physiol. 249, 387–398 (1975).

G. F. Cooper and J. G. Robson, "Application of Fourier analysis to the visibility of gratings," J. Physiol. 197, 551–566 (1968).

V. A. Movshon and D. J. Tolhurst, "On the response linearity of cells in the cat visual cortex," J. Physiol. 249, 56–57P (1975).

A. F. Dean, "The relationship between response amplitude and contrast for cat striate cortical neurones," J. Physiol. 318, 413–427 (1981).

Perception (1)

J. A. Movshon and C. Blakemore, "Orientation specificity and spatial selectivity in human vision," Perception 2, 53–60 (1973).

Proc. R. Soc. Lond. Ser. (1)

D. J. Tolhurst and I. D. Thompson, "On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat," Proc. R. Soc. Lond. Ser. B 213, 183–199 (1981).

Psychol. Rev. (1)

D. Regan, "Visual information channeling in normal and disordered vision," Psychol. Rev. 89, 407–444 (1982).

Vision Res. (1)

A. B. Watson and J. G. Robson, "Discrimination at threshold: labelled detectors in human vision," Vision Res. 21, 1115–1122 (1981).

Other (15)

H. R. Wilson and D. J. Gelb, "A modified line element theory for spatial frequency and width discrimination," J. Opt. Soc. Am. (to be published).

Presumably the most important among the neurons contributing to discrimination are those with the steepest slopes. However, it is not clear from published data on the spatial-frequency tuning of cortical cells whether the reason that neither symmetrically tuned neurons nor cells with steeper high- than low-frequency slopes seem to contribute importantly to discrimination is merely that these kinds of tuning curves are not associated with the steepest slopes.

J. Hirsch and R. Hylton, Department of Ophthalmology, Yale University, New Haven, Conn. 06520 (personal communication).

D. Regan, "Spatial frequency mechanisms in human vision: electrophysiological evidence," Vision Res. (to be published).

H. Wilson, Department of Biophysics and Theoretical Biology, University of Chicago, Chicago, Ill. 60637 (personal communication).

H. R. Wilson, D. K. McFarlane, and G. C. Phillips, "Spatial tuning of orientation selective units estimated by oblique masking," Vision Res. (to be published).

H. Wilson, Department of Biophysics and Theoretical Biology, University of Chicago, Chicago, Ill. 60637 (personal communication).

R. L. De Valois, D. G. Albrecht, and L. G. Thorell, "Cortical cells: bar and edge detectors, or spatial frequency filters?" in Frontiers in Visual Science, S. J. Cool and E. L. Smith, eds. (Springer, New York, 1978), pp. 544–556.

S1 and S2 would be the excitatory center/inhibitory surround type of neuron26 for pairs of dark lines and inhibitory center/excitatory surround26 for bright line pairs.

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

Caveats include the following: (1) 1(o take the bandwidth of the adaptation effect as the excitatory bandwidth of a channel implies that selective adaptation is a desensitization that is due to prolonged excitation, and there is some evidence that inhibition is involved.4 (2) In any case, the angular bandwidth of the threshold elevation is not necessarily the same as the angular bandwidth of a channel, since different channels may determine threshold before and after adaptation.5

This way of conceptually linking detection and discrimination is not restricted to spatial vision and has been applied to several visual modalities.19 In the context of color vision, Boynton20 discusses how the approach can be formulated either as a line element model, in which central neural processing has direct access to channel activities, or as an opponent process model, in which opponent mechanisms intervene between channels and subsequent processing. If they are both linear, the two formulations are equivalent. Historically, these two formulations of the relative activity approach have been influential and quite successful in color-vision research.20 More recently, this same concept has been invoked to account for acute discrimination between different orientations21 and acute discrimination of motion in depth.22

N. Graham, "Spatial frequency channels in human vision: detecting edges without edge detectors," in Visual Coding and Adaptability, C. S. Harris, ed. (Erlbaum, Hillsdale, N.J., 1980).

O. Braddick, F. W. Campbell and J. Atkinson, "Channels in vision: basic aspects," in Handbook of Sensory Physiology, H. W. Leibowitz and H.-L. Teuber, eds. (Springer, New York, 1978), Vol. 8.

D. H. Kelly and C. A. Burbeck, "Critical problems in spatial vision," in Critical Reuiews in Bioengineering (CRC, Boca Raton, Fla., 1983) (to be published).

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