Abstract

Monocular visual excitability was studied psychophysically by determining the threshold intensity for a small (40-min) brief (5-msec) “test” flash centered within a larger (3°) “conditioning” flash, both presented at 7° of arc in the temporal half-field. Interval between flashes was varied from −200 (test flash preceding onset of conditioning) to +1500 msec (test flash following), nine combinations of conditioning flash intensity and duration being used as parameters. For all combinations, threshold rose between −200 and −100 msec (before the start of the conditioning flash), reaching a maximum increment when the onsets of the two flashes were synchronized. When test was superimposed upon conditioning flash in time (positive intervals), threshold declined to approach an asymptote near the end of the latter flash. Following termination of the conditioning flash, threshold returned to resting level. The maximum rise in threshold, as well as the time taken to achieve an asymptote, varied with both intensity and duration of the conditioning flash. When a rough estimate of the photochemical contribution to these threshold changes was subtracted from the raw data, residual threshold increments were obtained and attributed to neural processes. On the basis of time course, it is suggested that these neural processes involve central (geniculocalcarine) as well as peripheral (neuro-retinal) factors.

© 1959 Optical Society of America

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References

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  1. S. Hecht, Physiol. Revs. 17, 239 (1937).
  2. G. Wald, Am. J. Ophthalmol. 40 (November, Pt. II), 18 (1955).
    [PubMed]
  3. R. Granit, Sensory Mechanisms of the Retina (Oxford University Press, New York, 1947).
  4. Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).
  5. H. K. Hartline and P. R. McDonald, J. Cellular Comp. Physiol. 30, 225 (1947).
    [Crossref]
  6. Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).
  7. W. Lohman, Z. Psychol. u. Physiol. der Sinnesorgane. Abt. II. Z. Sinnesphysiol. 41, 290 (1907).
  8. G. Wald and A. B. Clark, J. Gen. Physiol. 21, 93 (1937).
  9. H. K. Hartline and F. Ratliff, J. Gen. Physiol. 40, 357 (1957).
  10. S. W. Kuffler, J. Neurophysiol. 16, 37 (1953).
    [PubMed]
  11. R. J. Beitel, J. Gen. Psychol. 14, 31 (1936).
    [Crossref]
  12. A. Broca and D. Sulzer, J. physiol. et pathol. gén. 4, 632 (1902).
  13. G. A. Fry, Am. J. Physiol. 108, 701 (1934).
  14. G. A. Fry and S. H. Bartley, J. Exptl. Psychol. 19, 351 (1936).
    [Crossref]
  15. R. Granit and P. Harper, Am. J. Physiol. 95, 211 (1930).
  16. S. H. Bartley, J. Cellular Comp. Physiol. 8, 41 (1936).
    [Crossref]
  17. H. T. Chang, J. Neurophysiol. 13, 235 (1950).
  18. M. H. Clare and G. H. Bishop, EEG Clin. Neurophysiol. 4, 311 (1952).
    [Crossref]
  19. H. Gastaut and J. Hunter, EEG Clin. Neurophysiol. 2, 263 (1950).
    [Crossref]
  20. Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
    [Crossref]
  21. L. I. Malis and L. Krugcr, J. Neurophysiol. 19, 172 (1956).
    [PubMed]
  22. W. H. Marshall, J. Neurophysiol. 4, 277 (1949).
  23. Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).
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    [Crossref]
  25. W. S. Battersby, J. Exptl. Psychol. 42, 59 (1951).
    [Crossref]
  26. W. S. Battersby and M. B. Bender, J. Comp. and Physiol. Psychol. 51, 411 (1958).
    [Crossref]
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    [Crossref]
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    [Crossref] [PubMed]
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    [Crossref]
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    [PubMed]
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    [Crossref]
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    [Crossref]
  36. H. Gastaut, EEG Clin. Neurophysiol. 1, 205 (1949).
  37. M. Monnier, J. Neurophysiol. 15, 469 (1952).
    [PubMed]

1958 (1)

W. S. Battersby and M. B. Bender, J. Comp. and Physiol. Psychol. 51, 411 (1958).
[Crossref]

1957 (2)

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

H. K. Hartline and F. Ratliff, J. Gen. Physiol. 40, 357 (1957).

1956 (1)

L. I. Malis and L. Krugcr, J. Neurophysiol. 19, 172 (1956).
[PubMed]

1955 (4)

Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
[Crossref]

W. R. Bush, J. Opt. Soc. Am. 45, 1047 (1955).
[Crossref] [PubMed]

H. D. Baker, J. Opt. Soc. Am. 45, 839 (1955).
[Crossref] [PubMed]

G. Wald, Am. J. Ophthalmol. 40 (November, Pt. II), 18 (1955).
[PubMed]

1954 (1)

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

1953 (2)

R. M. Boynton and M. H. Triedman, J. Exptl. Psychol. 46, 125 (1953).
[Crossref]

S. W. Kuffler, J. Neurophysiol. 16, 37 (1953).
[PubMed]

1952 (2)

M. H. Clare and G. H. Bishop, EEG Clin. Neurophysiol. 4, 311 (1952).
[Crossref]

M. Monnier, J. Neurophysiol. 15, 469 (1952).
[PubMed]

1951 (1)

W. S. Battersby, J. Exptl. Psychol. 42, 59 (1951).
[Crossref]

1950 (2)

H. T. Chang, J. Neurophysiol. 13, 235 (1950).

H. Gastaut and J. Hunter, EEG Clin. Neurophysiol. 2, 263 (1950).
[Crossref]

1949 (2)

W. H. Marshall, J. Neurophysiol. 4, 277 (1949).

H. Gastaut, EEG Clin. Neurophysiol. 1, 205 (1949).

1947 (2)

B. H. Crawford, Proc. Roy. Soc. (London) B134, 283 (1947).
[Crossref]

H. K. Hartline and P. R. McDonald, J. Cellular Comp. Physiol. 30, 225 (1947).
[Crossref]

1943 (1)

Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).

1940 (2)

L. A. Riggs, J. Cellular Comp. Physiol. 15, 273 (1940).
[Crossref]

L. A. Riggs and C. H. Graham, J. Cellular Comp. Physiol. 16, 15 (1940).
[Crossref]

1939 (1)

Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).

1937 (3)

Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).

S. Hecht, Physiol. Revs. 17, 239 (1937).

G. Wald and A. B. Clark, J. Gen. Physiol. 21, 93 (1937).

1936 (4)

G. A. Fry and S. H. Bartley, J. Exptl. Psychol. 19, 351 (1936).
[Crossref]

S. H. Bartley, J. Cellular Comp. Physiol. 8, 41 (1936).
[Crossref]

R. J. Beitel, J. Gen. Psychol. 14, 31 (1936).
[Crossref]

G. H. Bishop, Cold Spring Harbor Symposium Quant. Biol. 4, 305 (1936).
[Crossref]

1934 (1)

G. A. Fry, Am. J. Physiol. 108, 701 (1934).

1930 (1)

R. Granit and P. Harper, Am. J. Physiol. 95, 211 (1930).

1907 (1)

W. Lohman, Z. Psychol. u. Physiol. der Sinnesorgane. Abt. II. Z. Sinnesphysiol. 41, 290 (1907).

1904 (1)

C. S. Sherrington, Brit. J. Psychol. 1, 26 (1904).

1902 (1)

A. Broca and D. Sulzer, J. physiol. et pathol. gén. 4, 632 (1902).

Ades,

Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).

Baker, H. D.

Bartley, S. H.

G. A. Fry and S. H. Bartley, J. Exptl. Psychol. 19, 351 (1936).
[Crossref]

S. H. Bartley, J. Cellular Comp. Physiol. 8, 41 (1936).
[Crossref]

Battersby, W. S.

W. S. Battersby and M. B. Bender, J. Comp. and Physiol. Psychol. 51, 411 (1958).
[Crossref]

W. S. Battersby, J. Exptl. Psychol. 42, 59 (1951).
[Crossref]

Beitel, R. J.

R. J. Beitel, J. Gen. Psychol. 14, 31 (1936).
[Crossref]

Bender, M. B.

W. S. Battersby and M. B. Bender, J. Comp. and Physiol. Psychol. 51, 411 (1958).
[Crossref]

Bishop, G. H.

M. H. Clare and G. H. Bishop, EEG Clin. Neurophysiol. 4, 311 (1952).
[Crossref]

G. H. Bishop, Cold Spring Harbor Symposium Quant. Biol. 4, 305 (1936).
[Crossref]

Boynton, R. M.

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

R. M. Boynton and M. H. Triedman, J. Exptl. Psychol. 46, 125 (1953).
[Crossref]

Broca, A.

A. Broca and D. Sulzer, J. physiol. et pathol. gén. 4, 632 (1902).

Bush, W. R.

Cahan,

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Chang, H. T.

H. T. Chang, J. Neurophysiol. 13, 235 (1950).

Chase,

Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).

Clare, M. H.

M. H. Clare and G. H. Bishop, EEG Clin. Neurophysiol. 4, 311 (1952).
[Crossref]

Clark, A. B.

G. Wald and A. B. Clark, J. Gen. Physiol. 21, 93 (1937).

Crawford, B. H.

B. H. Crawford, Proc. Roy. Soc. (London) B134, 283 (1947).
[Crossref]

Feinschil,

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Fry, G. A.

G. A. Fry and S. H. Bartley, J. Exptl. Psychol. 19, 351 (1936).
[Crossref]

G. A. Fry, Am. J. Physiol. 108, 701 (1934).

Gastaut, H.

H. Gastaut and J. Hunter, EEG Clin. Neurophysiol. 2, 263 (1950).
[Crossref]

H. Gastaut, EEG Clin. Neurophysiol. 1, 205 (1949).

Graham, C. H.

L. A. Riggs and C. H. Graham, J. Cellular Comp. Physiol. 16, 15 (1940).
[Crossref]

Granit,

Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).

Granit, R.

R. Granit and P. Harper, Am. J. Physiol. 95, 211 (1930).

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

Haig,

Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).

Harper, P.

R. Granit and P. Harper, Am. J. Physiol. 95, 211 (1930).

Hartline, H. K.

H. K. Hartline and F. Ratliff, J. Gen. Physiol. 40, 357 (1957).

H. K. Hartline and P. R. McDonald, J. Cellular Comp. Physiol. 30, 225 (1947).
[Crossref]

Hecht,

Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).

Hecht, S.

S. Hecht, Physiol. Revs. 17, 239 (1937).

Hunter, J.

H. Gastaut and J. Hunter, EEG Clin. Neurophysiol. 2, 263 (1950).
[Crossref]

Jasper,

Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
[Crossref]

Kandel, G.

King,

Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
[Crossref]

Krugcr, L.

L. I. Malis and L. Krugcr, J. Neurophysiol. 19, 172 (1956).
[PubMed]

Kuffler, S. W.

S. W. Kuffler, J. Neurophysiol. 16, 37 (1953).
[PubMed]

Lohman, W.

W. Lohman, Z. Psychol. u. Physiol. der Sinnesorgane. Abt. II. Z. Sinnesphysiol. 41, 290 (1907).

Malis, L. I.

L. I. Malis and L. Krugcr, J. Neurophysiol. 19, 172 (1956).
[PubMed]

Marshall,

Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).

Marshall, W. H.

W. H. Marshall, J. Neurophysiol. 4, 277 (1949).

McDonald, P. R.

H. K. Hartline and P. R. McDonald, J. Cellular Comp. Physiol. 30, 225 (1947).
[Crossref]

Monnier, M.

M. Monnier, J. Neurophysiol. 15, 469 (1952).
[PubMed]

Munsterhjelm,

Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).

Naidoff,

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Naquet,

Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
[Crossref]

Ratliff, F.

H. K. Hartline and F. Ratliff, J. Gen. Physiol. 40, 357 (1957).

Riggs, L. A.

L. A. Riggs, J. Cellular Comp. Physiol. 15, 273 (1940).
[Crossref]

L. A. Riggs and C. H. Graham, J. Cellular Comp. Physiol. 16, 15 (1940).
[Crossref]

Sherrington, C. S.

C. S. Sherrington, Brit. J. Psychol. 1, 26 (1904).

Sulzer, D.

A. Broca and D. Sulzer, J. physiol. et pathol. gén. 4, 632 (1902).

Talbot,

Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).

Triedman, M. H.

R. M. Boynton and M. H. Triedman, J. Exptl. Psychol. 46, 125 (1953).
[Crossref]

Wagman,

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Wald, G.

G. Wald, Am. J. Ophthalmol. 40 (November, Pt. II), 18 (1955).
[PubMed]

G. Wald and A. B. Clark, J. Gen. Physiol. 21, 93 (1937).

Waldman,

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Zewi,

Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).

Am. J. Ophthalmol. (2)

G. Wald, Am. J. Ophthalmol. 40 (November, Pt. II), 18 (1955).
[PubMed]

Wagman, Waldman, Naidoff, Feinschil, and Cahan, Am. J. Ophthalmol. 38, 60 (1954).
[PubMed]

Am. J. Physiol. (2)

R. Granit and P. Harper, Am. J. Physiol. 95, 211 (1930).

G. A. Fry, Am. J. Physiol. 108, 701 (1934).

Brit. J. Psychol. (1)

C. S. Sherrington, Brit. J. Psychol. 1, 26 (1904).

Cold Spring Harbor Symposium Quant. Biol. (1)

G. H. Bishop, Cold Spring Harbor Symposium Quant. Biol. 4, 305 (1936).
[Crossref]

EEG Clin. Neurophysiol. (4)

H. Gastaut, EEG Clin. Neurophysiol. 1, 205 (1949).

M. H. Clare and G. H. Bishop, EEG Clin. Neurophysiol. 4, 311 (1952).
[Crossref]

H. Gastaut and J. Hunter, EEG Clin. Neurophysiol. 2, 263 (1950).
[Crossref]

Jasper, Naquet, and King, EEG Clin. Neurophysiol. 7, 99 (1955).
[Crossref]

J. Cellular Comp. Physiol. (4)

H. K. Hartline and P. R. McDonald, J. Cellular Comp. Physiol. 30, 225 (1947).
[Crossref]

L. A. Riggs, J. Cellular Comp. Physiol. 15, 273 (1940).
[Crossref]

L. A. Riggs and C. H. Graham, J. Cellular Comp. Physiol. 16, 15 (1940).
[Crossref]

S. H. Bartley, J. Cellular Comp. Physiol. 8, 41 (1936).
[Crossref]

J. Comp. and Physiol. Psychol. (1)

W. S. Battersby and M. B. Bender, J. Comp. and Physiol. Psychol. 51, 411 (1958).
[Crossref]

J. Exptl. Psychol. (3)

W. S. Battersby, J. Exptl. Psychol. 42, 59 (1951).
[Crossref]

R. M. Boynton and M. H. Triedman, J. Exptl. Psychol. 46, 125 (1953).
[Crossref]

G. A. Fry and S. H. Bartley, J. Exptl. Psychol. 19, 351 (1936).
[Crossref]

J. Gen. Physiol. (3)

Hecht, Haig, and Chase, J. Gen. Physiol. 20, 831 (1937).

G. Wald and A. B. Clark, J. Gen. Physiol. 21, 93 (1937).

H. K. Hartline and F. Ratliff, J. Gen. Physiol. 40, 357 (1957).

J. Gen. Psychol. (1)

R. J. Beitel, J. Gen. Psychol. 14, 31 (1936).
[Crossref]

J. Neurophysiol. (6)

L. I. Malis and L. Krugcr, J. Neurophysiol. 19, 172 (1956).
[PubMed]

W. H. Marshall, J. Neurophysiol. 4, 277 (1949).

Marshall, Talbot, and Ades, J. Neurophysiol. 6, 1 (1943).

S. W. Kuffler, J. Neurophysiol. 16, 37 (1953).
[PubMed]

M. Monnier, J. Neurophysiol. 15, 469 (1952).
[PubMed]

H. T. Chang, J. Neurophysiol. 13, 235 (1950).

J. Opt. Soc. Am. (3)

J. Physiol. (1)

Granit, Munsterhjelm, and Zewi, J. Physiol. 96, 31 (1939).

J. physiol. et pathol. gén. (1)

A. Broca and D. Sulzer, J. physiol. et pathol. gén. 4, 632 (1902).

Physiol. Revs. (1)

S. Hecht, Physiol. Revs. 17, 239 (1937).

Proc. Roy. Soc. (London) (1)

B. H. Crawford, Proc. Roy. Soc. (London) B134, 283 (1947).
[Crossref]

Z. Psychol. u. Physiol. der Sinnesorgane. Abt. II. Z. Sinnesphysiol. (1)

W. Lohman, Z. Psychol. u. Physiol. der Sinnesorgane. Abt. II. Z. Sinnesphysiol. 41, 290 (1907).

Other (1)

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

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Figures (5)

F. 1
F. 1

Diagram of dual optical stimulator. With baffle B1 in front of P3, the two beams are superimposed for monocular stimulation of the right eye as described in text. With baffle B1 in front of combining prism (CP2), the beams remain independent, the right eye being stimulated by one, and the left eye by the other. Sketch of observer’s view is indicated in insert, along with the path of light rays emanating from the diagraphms. Not to scale.

F. 2
F. 2

Shutter control—dual optical stimulator. Flow diagram of electronic control circuits, plus detail of shutter mechanisms in insert. CR01, P1, and P2 indicate conventional amplifier-recording equipment, not used in present study. Sample records of light pulses as seen on monitor oscilloscope CRO2 given at the bottom. Three superimposed sweeps of the 5-msec test flash are presented to indicate reliability of shutter operation. See text for further details.

F. 3
F. 3

Schematic diagram illustrating the temporal relationships between test and conditioning flashes for five durations and three intensity levels of the latter. Intervals between flashes (delay) read algebraically from start of conditioning flash, and were so chosen for three durations of this flash (0.005, 0.050, 0.500 sec), that a determination could be made on the effects of this variable on threshold at fixed intervals of time.

F. 4
F. 4

Threshold in log10 mL along the ordinate at various intervals between test (Ft) and conditioning (Fc) flashes; negative intervals refer to condition where Ft precedes Fc in time. Three groups of curves are presented per observer, one for each of three intensity levels of Fc. At each intensity three curves are given, one for each of the three indicated durations of Fc. Thresholds measured at the end of Fc enclosed in squares. The range of resting threshold variability indicated by shaded horizontal bar at bottom. All points plotted were higher than or equal to resting threshold on day of testing. The dashed curve rising from 0 delay is the reciprocal of the dark adaptation process following cessation of the 500-msec. conditioning flash. It represents an approximation of the photochemical contribution to threshold change.

F. 5
F. 5

Estimated neural effect on threshold, obtained by subtracting presumed photochemical effect of light adaptation (rising dashed line) from data given in Fig. 4. Notation same as in Fig. 4.