Abstract

Electroretinograms and visually evoked cortical potentials, dependent on the occurence of spontaneous saccadic eye movements, were recorded from human observers. A computer system was used to isolate average-response waveforms that result from saccadic displacements of the retinal image. The responses were recorded as the observer fixed his eye on the center of a steady pattern of vertical stripes presented in maxwellian view. The responses depended on the spatial frequency of the pattern being viewed; in all cases, they were proportional to the extent of saccadic movement. The results are interpreted in terms of response additivity and the numbers of cones stimulated by the image displacement.

© 1974 Optical Society of America

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References

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  1. M. Alpern, in The Eye, Vol. 3, edited by H. Davson, (Academic, New York, 1962), Ch. 5.
  2. F. Ratliff and L. A. Riggs, J. Exper. Psychol. 40, 687 (1950).
    [Crossref]
  3. Handbook of Sensory Physiology, Vol. VII/4, edited by M. Alpern, D. Jameson, and L. M. Hurvich (Springer, Berlin, 1972), Ch. 12.
    [Crossref]
  4. L. Matin, in Ref. 3, Ch. 13.
  5. F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).
  6. F. Ratliff, J. Exper. Psychol. 43, 163 (1952).
    [Crossref]
  7. U. T. Keesey, J. Opt. Soc. Am. 50, 769 (1960).
    [Crossref] [PubMed]
  8. R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
    [Crossref] [PubMed]
  9. L. A. Riggs, F. Ratliff, J. Cornsweet, and T. N. Cornsweet, J. Opt. Soc. Am. 43, 495 (1953).
    [Crossref] [PubMed]
  10. R. W. Ditchburn and B. L. Ginsborg, Nature 170, 36 (1952).
    [Crossref] [PubMed]
  11. T. N. Cornsweet, J. Opt. Soc. Am. 46, 987 (1956).
    [Crossref] [PubMed]
  12. J. Nachmias, J. Opt. Soc. Am. 50, 569 (1960).
    [Crossref] [PubMed]
  13. R. J. Cunitz and R. M. Steinman, Vision Res. 9, 683 (1969).
    [Crossref] [PubMed]
  14. K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
    [Crossref] [PubMed]
  15. J. C. Armington, K. Gaarder, and A. M. L. Schick, J. Opt. Soc. Am. 57, 1534 (1967).
    [Crossref] [PubMed]
  16. E. D. Adrian, Mechanism of Nervous Action (Oxford U. P., Oxford, 1932).
  17. H. K. Hartline and J. Cell, Comp. Physiol. 5, 229 (1934).
  18. J. C. Armington, Behav. Res. Meth. Instru. 4(2), 61 (1972).
    [Crossref]
  19. L. E. Flamm, PhD thesis (Northeastern University, Boston, 1971).
  20. A. Skavenski and G. Timberlake, personal communication.
  21. K. Gaarder, Science 132, 471 (1960).
    [Crossref] [PubMed]
  22. J. C. Armington, J. Opt. Soc. Am. 49, 1169 (1959).
    [Crossref] [PubMed]
  23. J. C. Armington, Vision Res. 8, 263 (1968).
    [Crossref] [PubMed]
  24. H. J. M. Gerrits and A. J. H. Vendrik, Vision Res. 10, 1443 (1970).
    [Crossref] [PubMed]
  25. A. Troelstra and N. M. J. Schweitzer, J. Neurophysiol. 31, 588 (1968).
    [PubMed]
  26. L. H. van der Tweel, Annal. N. Y. Acad. Sci. 89, 829 (1961).
    [Crossref]
  27. E. P. Johnson, A. M. A. Arch. Ophthalmol. 60, 565 (1958).
    [Crossref]
  28. R. Granit, Sensory Mechanisms of the Retina (Hafner, New York, 1963).
  29. R. M. Boynton, J. Opt. Soc. Am. 43, 442 (1953).
    [Crossref] [PubMed]
  30. G. S. Brindley, J. Physiol. (Lond.) 134, 353 (1956).
  31. J. C. Armington, J. Opt. Soc. Am. 49, 1169 (1959).
    [Crossref] [PubMed]
  32. E. P. Johnson and N. R. Bartlett, J. Opt. Soc. Am. 46, 167 (1956).
    [Crossref] [PubMed]
  33. N. W. Perry and D. G. Childers, The Human Visual Evoked Response: Method and Theory (Thomas, Springfield, Ill., 1969), see pp. 46–48.
  34. E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).
  35. J. C. Armington, T. R. Corwin, and R. Marsetta, J. Opt. Soc. Am. 61, 1514 (1971).
    [Crossref] [PubMed]
  36. M. R. Harter and C. T. White, Vision Res. 8, 701 (1968).
    [Crossref] [PubMed]
  37. H. Spekreijse, Analysis of EEG Responses in Man Evoked by Sine Wave Modulated Light (Junk, The Hague, 1966).
  38. F. R. Ratliff, Mach Bands: Quantitative Studies on Neural Networks in the Retina (Holden-Day, San Francisco, 1965).
  39. J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
    [Crossref] [PubMed]
  40. This correction is taken from a study of alternating stimuli conducted by Armington, Corwin and Marsetta (Ref. 35). Figure 6 from Ref. 35 plots the relative amplitudes of the electroretinogram vs spatial frequency—other conditions being held constant. The amplitudes of the electroretinograms for the three stripe sizes plotted here were multiplied by factors inversely proportional to the attenuations found at those frequencies in Ref. 35.
  41. T. Oesterberg, Acta Ophthal. Suppl. 6, 1 (1935).
    [Crossref]

1973 (1)

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

1972 (1)

J. C. Armington, Behav. Res. Meth. Instru. 4(2), 61 (1972).
[Crossref]

1971 (1)

1970 (2)

J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
[Crossref] [PubMed]

H. J. M. Gerrits and A. J. H. Vendrik, Vision Res. 10, 1443 (1970).
[Crossref] [PubMed]

1969 (1)

R. J. Cunitz and R. M. Steinman, Vision Res. 9, 683 (1969).
[Crossref] [PubMed]

1968 (3)

A. Troelstra and N. M. J. Schweitzer, J. Neurophysiol. 31, 588 (1968).
[PubMed]

M. R. Harter and C. T. White, Vision Res. 8, 701 (1968).
[Crossref] [PubMed]

J. C. Armington, Vision Res. 8, 263 (1968).
[Crossref] [PubMed]

1967 (1)

1966 (1)

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).

1964 (1)

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

1961 (1)

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

1960 (3)

1959 (2)

1958 (1)

E. P. Johnson, A. M. A. Arch. Ophthalmol. 60, 565 (1958).
[Crossref]

1956 (3)

1953 (2)

1952 (2)

R. W. Ditchburn and B. L. Ginsborg, Nature 170, 36 (1952).
[Crossref] [PubMed]

F. Ratliff, J. Exper. Psychol. 43, 163 (1952).
[Crossref]

1950 (1)

F. Ratliff and L. A. Riggs, J. Exper. Psychol. 40, 687 (1950).
[Crossref]

1935 (1)

T. Oesterberg, Acta Ophthal. Suppl. 6, 1 (1935).
[Crossref]

1934 (1)

H. K. Hartline and J. Cell, Comp. Physiol. 5, 229 (1934).

1923 (1)

F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).

Adrian, E. D.

E. D. Adrian, Mechanism of Nervous Action (Oxford U. P., Oxford, 1932).

Alpern, M.

M. Alpern, in The Eye, Vol. 3, edited by H. Davson, (Academic, New York, 1962), Ch. 5.

Andersen, E. E.

F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).

Armington, J. C.

J. C. Armington, Behav. Res. Meth. Instru. 4(2), 61 (1972).
[Crossref]

J. C. Armington, T. R. Corwin, and R. Marsetta, J. Opt. Soc. Am. 61, 1514 (1971).
[Crossref] [PubMed]

J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
[Crossref] [PubMed]

J. C. Armington, Vision Res. 8, 263 (1968).
[Crossref] [PubMed]

J. C. Armington, K. Gaarder, and A. M. L. Schick, J. Opt. Soc. Am. 57, 1534 (1967).
[Crossref] [PubMed]

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

J. C. Armington, J. Opt. Soc. Am. 49, 1169 (1959).
[Crossref] [PubMed]

J. C. Armington, J. Opt. Soc. Am. 49, 1169 (1959).
[Crossref] [PubMed]

Averill, H. L.

F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).

Bartlett, N. R.

Boynton, R. M.

Brindley, G. S.

G. S. Brindley, J. Physiol. (Lond.) 134, 353 (1956).

Cell, J.

H. K. Hartline and J. Cell, Comp. Physiol. 5, 229 (1934).

Childers, D. G.

N. W. Perry and D. G. Childers, The Human Visual Evoked Response: Method and Theory (Thomas, Springfield, Ill., 1969), see pp. 46–48.

Cornsweet, J.

Cornsweet, T. N.

Corwin, T. R.

Cunitz, R. J.

R. J. Cunitz and R. M. Steinman, Vision Res. 9, 683 (1969).
[Crossref] [PubMed]

Ditchburn, R. W.

R. W. Ditchburn and B. L. Ginsborg, Nature 170, 36 (1952).
[Crossref] [PubMed]

Flamm, L. E.

L. E. Flamm, PhD thesis (Northeastern University, Boston, 1971).

Gaarder, K.

J. C. Armington, K. Gaarder, and A. M. L. Schick, J. Opt. Soc. Am. 57, 1534 (1967).
[Crossref] [PubMed]

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

K. Gaarder, Science 132, 471 (1960).
[Crossref] [PubMed]

Gerrits, H. J. M.

H. J. M. Gerrits and A. J. H. Vendrik, Vision Res. 10, 1443 (1970).
[Crossref] [PubMed]

Ginsborg, B. L.

R. W. Ditchburn and B. L. Ginsborg, Nature 170, 36 (1952).
[Crossref] [PubMed]

Graf, V.

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

Granit, R.

R. Granit, Sensory Mechanisms of the Retina (Hafner, New York, 1963).

Haddad, G. M.

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

Harter, M. R.

M. R. Harter and C. T. White, Vision Res. 8, 701 (1968).
[Crossref] [PubMed]

Hartline, H. K.

H. K. Hartline and J. Cell, Comp. Physiol. 5, 229 (1934).

Johnson, E. P.

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).

E. P. Johnson, A. M. A. Arch. Ophthalmol. 60, 565 (1958).
[Crossref]

E. P. Johnson and N. R. Bartlett, J. Opt. Soc. Am. 46, 167 (1956).
[Crossref] [PubMed]

Keesey, U. T.

Krauskopf, J.

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

Kropfl, W.

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

Marsetta, R.

J. C. Armington, T. R. Corwin, and R. Marsetta, J. Opt. Soc. Am. 61, 1514 (1971).
[Crossref] [PubMed]

J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
[Crossref] [PubMed]

Matin, L.

L. Matin, in Ref. 3, Ch. 13.

Nachmias, J.

Oesterberg, T.

T. Oesterberg, Acta Ophthal. Suppl. 6, 1 (1935).
[Crossref]

Perry, N. W.

N. W. Perry and D. G. Childers, The Human Visual Evoked Response: Method and Theory (Thomas, Springfield, Ill., 1969), see pp. 46–48.

Ratliff, F.

L. A. Riggs, F. Ratliff, J. Cornsweet, and T. N. Cornsweet, J. Opt. Soc. Am. 43, 495 (1953).
[Crossref] [PubMed]

F. Ratliff, J. Exper. Psychol. 43, 163 (1952).
[Crossref]

F. Ratliff and L. A. Riggs, J. Exper. Psychol. 40, 687 (1950).
[Crossref]

Ratliff, F. R.

F. R. Ratliff, Mach Bands: Quantitative Studies on Neural Networks in the Retina (Holden-Day, San Francisco, 1965).

Riggs, L. A.

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).

L. A. Riggs, F. Ratliff, J. Cornsweet, and T. N. Cornsweet, J. Opt. Soc. Am. 43, 495 (1953).
[Crossref] [PubMed]

F. Ratliff and L. A. Riggs, J. Exper. Psychol. 40, 687 (1950).
[Crossref]

Schick, A. M. L.

J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
[Crossref] [PubMed]

J. C. Armington, K. Gaarder, and A. M. L. Schick, J. Opt. Soc. Am. 57, 1534 (1967).
[Crossref] [PubMed]

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).

Schweitzer, N. M. J.

A. Troelstra and N. M. J. Schweitzer, J. Neurophysiol. 31, 588 (1968).
[PubMed]

Skavenski, A.

A. Skavenski and G. Timberlake, personal communication.

Skavenski, A. A.

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

Spekreijse, H.

H. Spekreijse, Analysis of EEG Responses in Man Evoked by Sine Wave Modulated Light (Junk, The Hague, 1966).

Steinman, R. M.

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

R. J. Cunitz and R. M. Steinman, Vision Res. 9, 683 (1969).
[Crossref] [PubMed]

Timberlake, G.

A. Skavenski and G. Timberlake, personal communication.

Troelstra, A.

A. Troelstra and N. M. J. Schweitzer, J. Neurophysiol. 31, 588 (1968).
[PubMed]

van der Tweel, L. H.

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

Vendrik, A. J. H.

H. J. M. Gerrits and A. J. H. Vendrik, Vision Res. 10, 1443 (1970).
[Crossref] [PubMed]

Weymouth, F. W.

F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).

White, C. T.

M. R. Harter and C. T. White, Vision Res. 8, 701 (1968).
[Crossref] [PubMed]

Wyman, D.

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

A. M. A. Arch. Ophthalmol. (1)

E. P. Johnson, A. M. A. Arch. Ophthalmol. 60, 565 (1958).
[Crossref]

Acta Ophthal. Suppl. (1)

T. Oesterberg, Acta Ophthal. Suppl. 6, 1 (1935).
[Crossref]

Am. J. Physiol. (1)

F. W. Weymouth, E. E. Andersen, and H. L. Averill, Am. J. Physiol. 63, 410 (1923).

Annal. N. Y. Acad. Sci. (1)

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

Behav. Res. Meth. Instru. (1)

J. C. Armington, Behav. Res. Meth. Instru. 4(2), 61 (1972).
[Crossref]

Comp. Physiol. (1)

H. K. Hartline and J. Cell, Comp. Physiol. 5, 229 (1934).

J. Exper. Psychol. (2)

F. Ratliff, J. Exper. Psychol. 43, 163 (1952).
[Crossref]

F. Ratliff and L. A. Riggs, J. Exper. Psychol. 40, 687 (1950).
[Crossref]

J. Neurophysiol. (1)

A. Troelstra and N. M. J. Schweitzer, J. Neurophysiol. 31, 588 (1968).
[PubMed]

J. Opt. Soc. Am. (10)

J. Physiol. (Lond.) (1)

G. S. Brindley, J. Physiol. (Lond.) 134, 353 (1956).

Nature (1)

R. W. Ditchburn and B. L. Ginsborg, Nature 170, 36 (1952).
[Crossref] [PubMed]

Science (3)

R. M. Steinman, G. M. Haddad, A. A. Skavenski, and D. Wyman, Science 181, 810 (1973).
[Crossref] [PubMed]

K. Gaarder, J. Krauskopf, V. Graf, W. Kropfl, and J. C. Armington, Science 146, 1481 (1964).
[Crossref] [PubMed]

K. Gaarder, Science 132, 471 (1960).
[Crossref] [PubMed]

Vision Res. (5)

J. C. Armington, Vision Res. 8, 263 (1968).
[Crossref] [PubMed]

H. J. M. Gerrits and A. J. H. Vendrik, Vision Res. 10, 1443 (1970).
[Crossref] [PubMed]

M. R. Harter and C. T. White, Vision Res. 8, 701 (1968).
[Crossref] [PubMed]

R. J. Cunitz and R. M. Steinman, Vision Res. 9, 683 (1969).
[Crossref] [PubMed]

J. C. Armington, R. Marsetta, and A. M. L. Schick, Vision Res. 10, 227 (1970).
[Crossref] [PubMed]

Vision Res. Suppl. (1)

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Vision Res. Suppl. 1, 75 (1966).

Other (11)

This correction is taken from a study of alternating stimuli conducted by Armington, Corwin and Marsetta (Ref. 35). Figure 6 from Ref. 35 plots the relative amplitudes of the electroretinogram vs spatial frequency—other conditions being held constant. The amplitudes of the electroretinograms for the three stripe sizes plotted here were multiplied by factors inversely proportional to the attenuations found at those frequencies in Ref. 35.

M. Alpern, in The Eye, Vol. 3, edited by H. Davson, (Academic, New York, 1962), Ch. 5.

E. D. Adrian, Mechanism of Nervous Action (Oxford U. P., Oxford, 1932).

L. E. Flamm, PhD thesis (Northeastern University, Boston, 1971).

A. Skavenski and G. Timberlake, personal communication.

Handbook of Sensory Physiology, Vol. VII/4, edited by M. Alpern, D. Jameson, and L. M. Hurvich (Springer, Berlin, 1972), Ch. 12.
[Crossref]

L. Matin, in Ref. 3, Ch. 13.

H. Spekreijse, Analysis of EEG Responses in Man Evoked by Sine Wave Modulated Light (Junk, The Hague, 1966).

F. R. Ratliff, Mach Bands: Quantitative Studies on Neural Networks in the Retina (Holden-Day, San Francisco, 1965).

N. W. Perry and D. G. Childers, The Human Visual Evoked Response: Method and Theory (Thomas, Springfield, Ill., 1969), see pp. 46–48.

R. Granit, Sensory Mechanisms of the Retina (Hafner, New York, 1963).

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Figures (7)

Fig. 1
Fig. 1

Examples of responses triggered by saccadic eye movements. Panel A. A comparison of responses produced by right and left movements. The electroretinogram is shown on the top, the average saccade in the middle and the visually evoked cortical potential at the bottom. Some alpha rhythm from the electroencephalogram that has not been completely averaged out is superimposed upon the right evoked potential. S, on the time scale, marks the point where the computer was triggered by the saccades. Note that the saccades begin before the trigger pulse, as explained in the text. Panel B shows the responses produced by a series of complex saccades to the left. Panel C illustrates the compound average-eye-movement waveform produced by a mixture of normal right saccades and left complex saccades. Panel D shows how the responses were measured. Observer JCA.

Fig. 2
Fig. 2

Amplitude histograms for saccadic eye movements produced while the observer viewed stripes of several angular subtenses. Each pair of histograms shows the frequency of eye movements in the right and left directions for the same experimental recording period. Observer JCA.

Fig. 3
Fig. 3

Comparison of electroretinograms and visually elicited cortical potentials produced by saccades of several amplitudes. A representative saccade waveform has been placed below each series of responses, to facilitate comparison. Note that different calibrations apply to the VECP and the ERG. Observer JCA.

Fig. 4
Fig. 4

Amplitude plots for the B wave. The amplitude of the B wave in microvolts is plotted against the angular rotation of the eye. The rotation, produced by saccadic movement, has been scaled in minutes of arc. Separate plots have been made for right and left rotations and for striped test patterns that subtended 6.6′, 13.2′, and 33′ of arc per stripe. Observer JCA.

Fig. 5
Fig. 5

Amplitude plots for the late-negative potential of the electroretinogram. This figure is similar to Fig. 4, except that the points describe a different component of the ERG. Observer JCA.

Fig. 6
Fig. 6

Amplitude plots for the VECP. These plots, constructed in the same manner as those in Fig. 4, relate the microvolt amplitude of the VECP to the angular rotation of the eye. Observer JCA.

Fig. 7
Fig. 7

The amplitude of the B wave as a function of the spatial frequency of the stimulus pattern. The upper plot shows the slope (the number of microvolts produced by a one minute rotation of the eye) of the B-wave-amplitude curves plotted against spatial frequency. Data from Fig. 4 for left and right movements have been combined. The lower curve shows the same data but the amplitudes have been corrected, as described in the text, for the reduction of amplitude of the ERG that occurs with closely patterned stimuli. Observer JCA.

Tables (1)

Tables Icon

Table I Correlations between the amplitudes of the B wave, the late-negative potential, and the VECP with saccadic amplitude for three subjects and several patterns. In a few cases (indicated by blank entries) no measurable response was obtained because of technical problems.