Abstract

Surface electrodes have been used for many years to pick up electrical activity that originates in the human eye or occipital cortex. In the earlier work, bright flashes of light were required to elicit responses that were large enough to measure against the ambient noise of the recording system. Recently, averaging techniques and other improvements have greatly increased the sensitivity of these methods. It is therefore no longer necessary to use the relatively meaningless light flashes, and responses can be obtained to visually significant aspects of the stimulus field. Of particular interest are specific effects due to image motion, color differences, and binocular fusion and rivalry. By the use of standard contact-lens and EEG electrodes, information on the processing of the stimulus pattern is obtained at both retinal and occipital levels of the human visual system. Microelectrode experiments with animal subjects have likewise shown that single flashes of light are of minor importance as visual stimuli, and that more significant aspects of visual patterns are particularly effective in arousing the responses of visual cells at all levels. Hence this evidence can be used as a guide to the interpretation of the human results, since these are necessarily obtained from electrodes that are remote from the sites at which they are generated. In short, the traditional methods of psychophysics are still used to provide definitive data on human vision. But the neural basis for these data is most clearly provided by microelectrode studies on the visual cells of experimental animals, together with such remotely recorded human action potentials as can be obtained from the retina and cortex.

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  4. R. Granit, Sensory Mechanisms of the Retina (Oxford University Press, New York, 1947).
  5. T. Tomita, J. Opt. Soc. Am. 53, 49 (1963); K. T. Brown, Vision Res. 8, 633 (1968).
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  13. R. Cooper, A. L. Winter, and W. G. Walter, Electroenceph. Clin. Neurophysiol. 18, 217 (1965); S. S. Fox and J. H. O'Brien, Science 147, 888 (1965); M. Rayport, H. G. Vaughan, and C. L. Rosengart, Electroenceph. Clin. Neurophysiol. 17, 608 (1964).
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  30. E. G. Gross, H. G. Vaughan, Jr., and E. Valenstein, Electroenceph. Clin. Neurophysiol. 22, 204 (1967).
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  32. J. H. Jacobson, Clinical Electroretinography (Charles C Thomas, Springfield, Ill., 1961).
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Adrian, E. D.

E. D. Adrian, Nature 154, 361 (1944).

E. D. Adrian, J. Physiol. (London) 105, 24 (1946); E. Dodt, Nature 168, 738 (1951); H. Bornschein and G. Schubert, Z. Biol. 106, 229 (1953).

E. D. Adrian, J. Physiol. (London) 104, 84 (1945); R. Granit, Receptors and Sensory Perception (Yale University Press, New Haven, 1955).

Beeler, Jr., G. W.

D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Electroenceph. Clin. Neurophysiol. 22, 136 (1967).

Beinhocker, G. D.

R. M. Copenhaver and G. D. Beinhocker, Invest. Ophthalmol. 2, 393 (1963); R. G. DeVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968); W. J. Rietveld, W. E. M. Tordoir, and J. W. Duyff, Acta Physiol. Pharmacol. Neerl. 13, 330 (1965).

Bergamasco, B.

L. Bergamini and B. Bergamasco, Cortical Evoked Potentials in Man (Charles C. Thomas, Springfield, Ill., 1967); R. M. Harter and C. T. White, Vision Res. 8, 701 (1968); W. J. Rietveld, W. E. M. Tordoir, J. R. B. Hagenouw, J. A. Lubbers, and A. C. Spoor, Acta Physiol. Pharmacol. Neerl. 14, 259 (1967); R. Spehlmann, Electroenceph. Clin. Neurophysiol. 19, 560 (1965); H. Spekreijse, Analysis of E. E. G. Responses in Man, thesis, University of Amsterdam (1966).

Bergamini, L.

L. Bergamini and B. Bergamasco, Cortical Evoked Potentials in Man (Charles C. Thomas, Springfield, Ill., 1967); R. M. Harter and C. T. White, Vision Res. 8, 701 (1968); W. J. Rietveld, W. E. M. Tordoir, J. R. B. Hagenouw, J. A. Lubbers, and A. C. Spoor, Acta Physiol. Pharmacol. Neerl. 14, 259 (1967); R. Spehlmann, Electroenceph. Clin. Neurophysiol. 19, 560 (1965); H. Spekreijse, Analysis of E. E. G. Responses in Man, thesis, University of Amsterdam (1966).

Berry, R. N.

L. A. Riggs, R. N. Berry, and M. Wayner, J. Opt. Soc. Am. 39, 427 (1949); J. C. Armington, D. I. Tepas, W. J. Kropfl, and W. H. Hengst, J. Opt. Soc. Am. 51, 877 (1961).

Boynton, R. M.

R. M. Boynton and L. A. Riggs, J. Exptl. Psychol. 42, 217 (1951); R. Boynton, J. Opt. Soc. Am. 43, 442 (1953).

Bremer, F.

F. Bremer, in Sensory Communications, W. A. Rosenblith, Ed. (John Wiley & Sons, Inc., New York, 1961); O. D. Creutzfeld, S. Watanabe, and H. D. Lux, Electroenceph. Clin. Neurophysiol. 20, 1 (1966).

Brown, J. L.

J. L. Brown, Ref. 9, p. 39.

Brown, K. T.

K. T. Brown and M. Murakami, Nature 201, 626 (1964).

Burkhardt, D. A.

D. A. Burkhardt and L. A. Riggs, Vision Res. 7, 453 (1967).

Chinn, H. I.

W. Noell and H. I. Chinn, Am. J. Physiol. 161, 573 (1950).

Cobb, W. A.

W. A. Cobb, H. B. Morton, and G. Ettlinger, Nature 216, 1123 (1967).

Cone, R. A.

R. A. Cone, Nature 204, 736 (1964).

Cooper, R.

R. Cooper, A. L. Winter, and W. G. Walter, Electroenceph. Clin. Neurophysiol. 18, 217 (1965); S. S. Fox and J. H. O'Brien, Science 147, 888 (1965); M. Rayport, H. G. Vaughan, and C. L. Rosengart, Electroenceph. Clin. Neurophysiol. 17, 608 (1964).

Copenhaver, R. M.

R. M. Copenhaver and G. D. Beinhocker, Invest. Ophthalmol. 2, 393 (1963); R. G. DeVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968); W. J. Rietveld, W. E. M. Tordoir, and J. W. Duyff, Acta Physiol. Pharmacol. Neerl. 13, 330 (1965).

Cornsweet, T. N.

E. P. Johnson and T. N. Cornsweet, Nature 174, 614 (1954); J. Armington, J. Opt. Soc. Am. 45, 1058 (1955).

DeValois, R. L.

R. L. DeValois, J. Gen. Physiol. 43, Suppl., Part 2, 115 (1960).

Elul, R.

R. Elul, in Data Acquisition and Processing in Biology and Medicine (Pergamon Press, Inc., New York, 1968), Vol. V, p. 93.

Ettlinger, G.

W. A. Cobb, H. B. Morton, and G. Ettlinger, Nature 216, 1123 (1967).

Fender, D. H.

D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Electroenceph. Clin. Neurophysiol. 22, 136 (1967).

D. Lehmann and D. H. Fender, Electroenceph. Clin. Neurophysiol. 24, 542 (1968); D. M. MacKay, Nature 217, 81 (1968); D. M. MacKay and W. J. Rietveld, Nature 217, 677 (1968).

Galloway, N. R.

N. R. Galloway, Brit. J. Ophthalmol. 51, 261 (1967); D. Yonemura and K. Kawasaki, Jap. J. Physiol. 17, 235 (1967).

Goldring, S.

Y. Karahashi and S. Goldring, Electroenceph. Clin. Neurophysiol. 20, 600 (1966).

Gouras, P.

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

Granit, R.

R. Granit, in The Eye, H. Davson, Ed. (Academic Press Inc., New York, 1962), Vol. II, p. 537.

R. Granit, Sensory Mechanisms of the Retina (Oxford University Press, New York, 1947).

Gross, E. G.

E. G. Gross, H. G. Vaughan, Jr., and E. Valenstein, Electroenceph. Clin. Neurophysiol. 22, 204 (1967).

Gunkel, R. D.

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

Haider, M.

M. Haider, P. Spong, and D. B. Lindsley, Science 145, 180 (1964); P. Spong, M. Haider, and D. B. Lindsley, Science 148, 395 (1965).

Hartline, H. K.

H. K. Hartline, Am. J. Physiol. 73, 600 (1925).

Hubel, D. H.

D. H. Hubel and T. N. Wiesel, J. Physiol. (London) 195, 215 (1968).

Jacobson, J. H.

J. H. Jacobson, Clinical Electroretinography (Charles C Thomas, Springfield, Ill., 1961).

Johnson, E. P.

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, J. Opt. Soc. Am. 56, 1621 (1966).

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Clinical Electroretinography (Suppl. to Vision Res.) (1966), p. 75.

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, Science 144, 567 (1964).

E. P. Johnson and T. N. Cornsweet, Nature 174, 614 (1954); J. Armington, J. Opt. Soc. Am. 45, 1058 (1955).

Karahashi, Y.

Y. Karahashi and S. Goldring, Electroenceph. Clin. Neurophysiol. 20, 600 (1966).

Karpe, G.

G. Karpe, Acta Ophthalmol. (Copenhagen) Suppl. 24, 1 (1945).

Kohlrausch, A.

A. Kohlrausch, in Handbuch der normalen und pathologischen Physiologie, A. Bethe, G. von Bergmann, G. Embden, and A. Ellinger, Eds. (Springer Verlag, Berlin, 1931), Vol. XII, Sec. 2, p. 1394.

Lehmann, D.

D. Lehmann and D. H. Fender, Electroenceph. Clin. Neurophysiol. 24, 542 (1968); D. M. MacKay, Nature 217, 81 (1968); D. M. MacKay and W. J. Rietveld, Nature 217, 677 (1968).

D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Electroenceph. Clin. Neurophysiol. 22, 136 (1967).

Lindsley, D. B.

M. Haider, P. Spong, and D. B. Lindsley, Science 145, 180 (1964); P. Spong, M. Haider, and D. B. Lindsley, Science 148, 395 (1965).

Morton, H. B.

W. A. Cobb, H. B. Morton, and G. Ettlinger, Nature 216, 1123 (1967).

Murakami, M.

K. T. Brown and M. Murakami, Nature 201, 626 (1964).

Noell, W.

W. Noell and H. I. Chinn, Am. J. Physiol. 161, 573 (1950).

Riggs, L. A.

L. A. Riggs, in Vision and Visual Perception, C. H. Graham, Ed. (John Wiley & Sons, Inc., New York, 1965), p. 81.

L. A. Riggs, Proc. Soc. Exptl. Biol. Med. 48, 204 (1941).

L. A. Riggs, R. N. Berry, and M. Wayner, J. Opt. Soc. Am. 39, 427 (1949); J. C. Armington, D. I. Tepas, W. J. Kropfl, and W. H. Hengst, J. Opt. Soc. Am. 51, 877 (1961).

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, Science 144, 567 (1964).

L. A. Riggs, Am. J. Ophthalmol. 38, 70 (1954).

R. M. Boynton and L. A. Riggs, J. Exptl. Psychol. 42, 217 (1951); R. Boynton, J. Opt. Soc. Am. 43, 442 (1953).

L. A. Riggs and P. Whittle, Vision Res. 7, 441 (1967); D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Nature 215, 204 (1967).

L. A. Riggs and C. E. Sternheim, J. Opt. Soc. Am. 59, 635 (1969).

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, J. Opt. Soc. Am. 56, 1621 (1966).

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Clinical Electroretinography (Suppl. to Vision Res.) (1966), p. 75.

C. E. Sternheim and L. A. Riggs, Vision Res. 8, 25 (1968).

D. A. Burkhardt and L. A. Riggs, Vision Res. 7, 453 (1967).

Schick, A. M. L.

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Clinical Electroretinography (Suppl. to Vision Res.) (1966), p. 75.

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, J. Opt. Soc. Am. 56, 1621 (1966).

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, Science 144, 567 (1964).

Siegfried, J. B.

J. B. Siegfried, Spectral Sensitivity of the Human Visual System as Measured from, Both Evoked Cortical Potentials and Psychophysical Responses, Ph.D. dissertation, Brown University (1967).

Spong, P.

M. Haider, P. Spong, and D. B. Lindsley, Science 145, 180 (1964); P. Spong, M. Haider, and D. B. Lindsley, Science 148, 395 (1965).

Sternheim, C. E.

C. E. Sternheim and L. A. Riggs, Vision Res. 8, 25 (1968).

L. A. Riggs and C. E. Sternheim, J. Opt. Soc. Am. 59, 635 (1969).

Tomita, T.

T. Tomita, J. Opt. Soc. Am. 53, 49 (1963); K. T. Brown, Vision Res. 8, 633 (1968).

Valenstein, E.

E. G. Gross, H. G. Vaughan, Jr., and E. Valenstein, Electroenceph. Clin. Neurophysiol. 22, 204 (1967).

Vaughan, Jr., H. G.

E. G. Gross, H. G. Vaughan, Jr., and E. Valenstein, Electroenceph. Clin. Neurophysiol. 22, 204 (1967).

Wald, G.

G. Wald, Science 101, 653 (1945).

Walter, W. G.

R. Cooper, A. L. Winter, and W. G. Walter, Electroenceph. Clin. Neurophysiol. 18, 217 (1965); S. S. Fox and J. H. O'Brien, Science 147, 888 (1965); M. Rayport, H. G. Vaughan, and C. L. Rosengart, Electroenceph. Clin. Neurophysiol. 17, 608 (1964).

Wayner, M.

L. A. Riggs, R. N. Berry, and M. Wayner, J. Opt. Soc. Am. 39, 427 (1949); J. C. Armington, D. I. Tepas, W. J. Kropfl, and W. H. Hengst, J. Opt. Soc. Am. 51, 877 (1961).

Whittle, P.

L. A. Riggs and P. Whittle, Vision Res. 7, 441 (1967); D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Nature 215, 204 (1967).

Wiesel, T. N.

D. H. Hubel and T. N. Wiesel, J. Physiol. (London) 195, 215 (1968).

Winter, A. L.

R. Cooper, A. L. Winter, and W. G. Walter, Electroenceph. Clin. Neurophysiol. 18, 217 (1965); S. S. Fox and J. H. O'Brien, Science 147, 888 (1965); M. Rayport, H. G. Vaughan, and C. L. Rosengart, Electroenceph. Clin. Neurophysiol. 17, 608 (1964).

Other (44)

A. Kohlrausch, in Handbuch der normalen und pathologischen Physiologie, A. Bethe, G. von Bergmann, G. Embden, and A. Ellinger, Eds. (Springer Verlag, Berlin, 1931), Vol. XII, Sec. 2, p. 1394.

H. K. Hartline, Am. J. Physiol. 73, 600 (1925).

L. A. Riggs, Proc. Soc. Exptl. Biol. Med. 48, 204 (1941).

R. Granit, Sensory Mechanisms of the Retina (Oxford University Press, New York, 1947).

T. Tomita, J. Opt. Soc. Am. 53, 49 (1963); K. T. Brown, Vision Res. 8, 633 (1968).

W. Noell and H. I. Chinn, Am. J. Physiol. 161, 573 (1950).

R. Granit, in The Eye, H. Davson, Ed. (Academic Press Inc., New York, 1962), Vol. II, p. 537.

G. Karpe, Acta Ophthalmol. (Copenhagen) Suppl. 24, 1 (1945).

L. A. Riggs, in Vision and Visual Perception, C. H. Graham, Ed. (John Wiley & Sons, Inc., New York, 1965), p. 81.

F. Bremer, in Sensory Communications, W. A. Rosenblith, Ed. (John Wiley & Sons, Inc., New York, 1961); O. D. Creutzfeld, S. Watanabe, and H. D. Lux, Electroenceph. Clin. Neurophysiol. 20, 1 (1966).

R. Elul, in Data Acquisition and Processing in Biology and Medicine (Pergamon Press, Inc., New York, 1968), Vol. V, p. 93.

Y. Karahashi and S. Goldring, Electroenceph. Clin. Neurophysiol. 20, 600 (1966).

R. Cooper, A. L. Winter, and W. G. Walter, Electroenceph. Clin. Neurophysiol. 18, 217 (1965); S. S. Fox and J. H. O'Brien, Science 147, 888 (1965); M. Rayport, H. G. Vaughan, and C. L. Rosengart, Electroenceph. Clin. Neurophysiol. 17, 608 (1964).

M. Haider, P. Spong, and D. B. Lindsley, Science 145, 180 (1964); P. Spong, M. Haider, and D. B. Lindsley, Science 148, 395 (1965).

K. T. Brown and M. Murakami, Nature 201, 626 (1964).

R. A. Cone, Nature 204, 736 (1964).

N. R. Galloway, Brit. J. Ophthalmol. 51, 261 (1967); D. Yonemura and K. Kawasaki, Jap. J. Physiol. 17, 235 (1967).

E. D. Adrian, Nature 154, 361 (1944).

L. A. Riggs, R. N. Berry, and M. Wayner, J. Opt. Soc. Am. 39, 427 (1949); J. C. Armington, D. I. Tepas, W. J. Kropfl, and W. H. Hengst, J. Opt. Soc. Am. 51, 877 (1961).

E. D. Adrian, J. Physiol. (London) 105, 24 (1946); E. Dodt, Nature 168, 738 (1951); H. Bornschein and G. Schubert, Z. Biol. 106, 229 (1953).

E. P. Johnson and T. N. Cornsweet, Nature 174, 614 (1954); J. Armington, J. Opt. Soc. Am. 45, 1058 (1955).

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, Science 144, 567 (1964).

E. P. Johnson, L. A. Riggs, and A. M. L. Schick, Clinical Electroretinography (Suppl. to Vision Res.) (1966), p. 75.

L. A. Riggs, E. P. Johnson, and A. M. L. Schick, J. Opt. Soc. Am. 56, 1621 (1966).

C. E. Sternheim and L. A. Riggs, Vision Res. 8, 25 (1968).

J. L. Brown, Ref. 9, p. 39.

R. M. Copenhaver and G. D. Beinhocker, Invest. Ophthalmol. 2, 393 (1963); R. G. DeVoe, H. Ripps, and H. G. Vaughan, Jr., Vision Res. 8, 135 (1968); W. J. Rietveld, W. E. M. Tordoir, and J. W. Duyff, Acta Physiol. Pharmacol. Neerl. 13, 330 (1965).

L. Bergamini and B. Bergamasco, Cortical Evoked Potentials in Man (Charles C. Thomas, Springfield, Ill., 1967); R. M. Harter and C. T. White, Vision Res. 8, 701 (1968); W. J. Rietveld, W. E. M. Tordoir, J. R. B. Hagenouw, J. A. Lubbers, and A. C. Spoor, Acta Physiol. Pharmacol. Neerl. 14, 259 (1967); R. Spehlmann, Electroenceph. Clin. Neurophysiol. 19, 560 (1965); H. Spekreijse, Analysis of E. E. G. Responses in Man, thesis, University of Amsterdam (1966).

D. A. Burkhardt and L. A. Riggs, Vision Res. 7, 453 (1967).

E. G. Gross, H. G. Vaughan, Jr., and E. Valenstein, Electroenceph. Clin. Neurophysiol. 22, 204 (1967).

W. A. Cobb, H. B. Morton, and G. Ettlinger, Nature 216, 1123 (1967).

J. H. Jacobson, Clinical Electroretinography (Charles C Thomas, Springfield, Ill., 1961).

L. A. Riggs, Am. J. Ophthalmol. 38, 70 (1954).

R. M. Boynton and L. A. Riggs, J. Exptl. Psychol. 42, 217 (1951); R. Boynton, J. Opt. Soc. Am. 43, 442 (1953).

E. D. Adrian, J. Physiol. (London) 104, 84 (1945); R. Granit, Receptors and Sensory Perception (Yale University Press, New Haven, 1955).

G. Wald, Science 101, 653 (1945).

J. B. Siegfried, Spectral Sensitivity of the Human Visual System as Measured from, Both Evoked Cortical Potentials and Psychophysical Responses, Ph.D. dissertation, Brown University (1967).

L. A. Riggs and C. E. Sternheim, J. Opt. Soc. Am. 59, 635 (1969).

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

R. L. DeValois, J. Gen. Physiol. 43, Suppl., Part 2, 115 (1960).

D. H. Hubel and T. N. Wiesel, J. Physiol. (London) 195, 215 (1968).

L. A. Riggs and P. Whittle, Vision Res. 7, 441 (1967); D. Lehmann, G. W. Beeler, Jr., and D. H. Fender, Nature 215, 204 (1967).

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