Abstract

A recently discovered component of the human electroretinogram (the electrical response wave resulting from stimulation by light) is recorded with an electrode mounted in a contact lens. Although previous investigators have thought it to be a photopic response, this component of the ERG is found to have a spectral sensitivity function that is maximal at approximately 630 mμ and is not similar to either a photopic or a scotopic luminosity function. This component is absent in the eyes of protanopes (red color blind) but present in eyes characterized by other forms of color deficiency. It increases rapidly with dark adaptation and is greatest after approximately one minute in darkness. It is accompanied by several new and previously unreported waves which also make their appearance during the early stages of dark adaptation. This research provides direct physiological evidence for a peripheral locus of color blindness and for a specific red color mechanism.

© 1952 Optical Society of America

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References

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  1. E. D. Adrian, J. Physiol. 104, 84 (1945).and J. Physiol. 105, 24 (1946).
  2. Riggs, Berry, and Wayner, J. Opt. Soc. Am. 39, 427 (1949).
    [Crossref] [PubMed]
  3. E. P. Johnson, J. Exptl. Psychol. 39, 597 (1949).
    [Crossref]
  4. K. Motokawa and T. Mita, Tohoku J. Exp. Med. 42, 114 (1942).
    [Crossref]
  5. L. A. Riggs, Proc. Soc. Exptl. Biol. Med. 48, 204 (1941).
    [Crossref]
  6. W. S. Stiles and T. Smith, Proc. Phys. Soc. (London) 56, 251 (1944).
    [Crossref]
  7. F. H. G. Pitt, Medical Research Council, Report of the Committee on the Physiology of Vision, XIV, (1935).
  8. L. L. Sloan, Psychol. Monogr. 38, 87 (1928).
  9. C. H. Graham and Y. Hsia, Proc. Natl. Acad. Sci. 38, 80 (1952).
    [Crossref]
  10. R. Granit, Oxford University Press, London, (1947).
  11. W. S. Stiles, Ned. T. Natuurk. 15, 125 (1949).

1952 (1)

C. H. Graham and Y. Hsia, Proc. Natl. Acad. Sci. 38, 80 (1952).
[Crossref]

1949 (3)

W. S. Stiles, Ned. T. Natuurk. 15, 125 (1949).

Riggs, Berry, and Wayner, J. Opt. Soc. Am. 39, 427 (1949).
[Crossref] [PubMed]

E. P. Johnson, J. Exptl. Psychol. 39, 597 (1949).
[Crossref]

1945 (1)

E. D. Adrian, J. Physiol. 104, 84 (1945).and J. Physiol. 105, 24 (1946).

1944 (1)

W. S. Stiles and T. Smith, Proc. Phys. Soc. (London) 56, 251 (1944).
[Crossref]

1942 (1)

K. Motokawa and T. Mita, Tohoku J. Exp. Med. 42, 114 (1942).
[Crossref]

1941 (1)

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

1928 (1)

L. L. Sloan, Psychol. Monogr. 38, 87 (1928).

Adrian, E. D.

E. D. Adrian, J. Physiol. 104, 84 (1945).and J. Physiol. 105, 24 (1946).

Berry,

Graham, C. H.

C. H. Graham and Y. Hsia, Proc. Natl. Acad. Sci. 38, 80 (1952).
[Crossref]

Granit, R.

R. Granit, Oxford University Press, London, (1947).

Hsia, Y.

C. H. Graham and Y. Hsia, Proc. Natl. Acad. Sci. 38, 80 (1952).
[Crossref]

Johnson, E. P.

E. P. Johnson, J. Exptl. Psychol. 39, 597 (1949).
[Crossref]

Mita, T.

K. Motokawa and T. Mita, Tohoku J. Exp. Med. 42, 114 (1942).
[Crossref]

Motokawa, K.

K. Motokawa and T. Mita, Tohoku J. Exp. Med. 42, 114 (1942).
[Crossref]

Pitt, F. H. G.

F. H. G. Pitt, Medical Research Council, Report of the Committee on the Physiology of Vision, XIV, (1935).

Riggs,

Riggs, L. A.

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

Sloan, L. L.

L. L. Sloan, Psychol. Monogr. 38, 87 (1928).

Smith, T.

W. S. Stiles and T. Smith, Proc. Phys. Soc. (London) 56, 251 (1944).
[Crossref]

Stiles, W. S.

W. S. Stiles, Ned. T. Natuurk. 15, 125 (1949).

W. S. Stiles and T. Smith, Proc. Phys. Soc. (London) 56, 251 (1944).
[Crossref]

Wayner,

J. Exptl. Psychol. (1)

E. P. Johnson, J. Exptl. Psychol. 39, 597 (1949).
[Crossref]

J. Opt. Soc. Am. (1)

J. Physiol. (1)

E. D. Adrian, J. Physiol. 104, 84 (1945).and J. Physiol. 105, 24 (1946).

Ned. T. Natuurk. (1)

W. S. Stiles, Ned. T. Natuurk. 15, 125 (1949).

Proc. Natl. Acad. Sci. (1)

C. H. Graham and Y. Hsia, Proc. Natl. Acad. Sci. 38, 80 (1952).
[Crossref]

Proc. Phys. Soc. (London) (1)

W. S. Stiles and T. Smith, Proc. Phys. Soc. (London) 56, 251 (1944).
[Crossref]

Proc. Soc. Exptl. Biol. Med. (1)

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

Psychol. Monogr. (1)

L. L. Sloan, Psychol. Monogr. 38, 87 (1928).

Tohoku J. Exp. Med. (1)

K. Motokawa and T. Mita, Tohoku J. Exp. Med. 42, 114 (1942).
[Crossref]

Other (2)

F. H. G. Pitt, Medical Research Council, Report of the Committee on the Physiology of Vision, XIV, (1935).

R. Granit, Oxford University Press, London, (1947).

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Figures (13)

Fig. 1
Fig. 1

Diagram of the stimulation system. The letters point out the following: P, subject’s pupil; SURR, adaptation screen; M1 and M2, unsilvered mirrors reflecting light from the fixation point into the main optical path; S motor driven disk shutter; A, diaphragm stop; F, optical filters; and LAMP, main tungsten stimulus scource. The shield prevents alternating current interference from reaching the subject’s electrodes. The stimulus indication beam makes a trace on the photographic record whenever a stimulus is administered (adapted from diagram by Johnson, reference 3).

Fig. 2
Fig. 2

Block diagram of the principal units of the recording equipment. Arrows indicate direction of travel of retinal potentials. MG is a monitoring galvanometer used to maintain the direct-coupled amplifier in a balanced condition. The oscilloscope is used in such a manner that the experimenter may simultaneously observe and record filtered responses. Both unfiltered (DC) and filtered responses are simultaneously recorded on the same photographic tape.

Fig. 3
Fig. 3

Variations in the ERG as a result of changes in stimulus wavelength. Subject JW. Peaks of transmission of filter in mμ: B 640, C 630, D 616, E 573, F 545, G 533, H 510, I 467.

Fig. 4
Fig. 4

Typical X waves. To the left are shown X waves from two subjects. Tracing on the right illustrates the response of the capacitance-coupled amplification system to rectangular calibration pulses.

Fig. 5
Fig. 5

Response sensitivity in relation to photopic and scotopic luminosity. The X wave appears to be related to only the red extremity of the photopic luminosity function. A scotopic response rather than the X wave is recorded throughout most of the spectrum.

Fig. 6
Fig. 6

Responses of normal and red blind subjects to red and white light. The two classes of subject cannot be distinguished with white stimuli, but protanopes show reduced activity with red. Photographs are from the filtered record; heavy line on PD’s response to white light is the direct-coupled record which in this case ran over the filtered line.

Fig. 7
Fig. 7

Response amplitude as a function of stimulus intensity. Stimulus intensity increases to the right; numbers refer to the density of neutral filtering used. The question mark beside one circle indicates that no detectable response was produced by this subject; the circle is merely drawn at the noise level of his record. With white light a log stimulus intensity value of 0.0 is 5 log units above 1 ft-L. (The method of making this measurement is outlined in footnote §.)

Fig. 8
Fig. 8

Comparisons of response sensitivity with the scotopic luminosity function in normal and color blind eyes. Only protanopes (red blind) show a lack of red X sensitivity.

Fig. 9
Fig. 9

A schematic drawing (not to scale) of the apparatus used to provide a means of bright light adaptation. A, final lens of main stimulation system; B, unsilvered mirror; C, position of observer’s eye; D, lens; E, neutral filter; F, heat filter; G, tungsten filament source.

Fig. 10
Fig. 10

Initial course of X wave recovery. For each observer curves were obtained with stimuli 0.0, 0.3, 0.6, and 1.0 log unit below maximum intensity. (The uppermost curve is for an intensity of 0.0 log unit.)

Fig. 11
Fig. 11

Plots of response recovery during dark adaptation. Dashed line is for the X wave (filtered record); solid line for the B wave (direct-coupled record). Upper four rows were obtained using the FR filter and neutral filtering of 0.0, 0.3, 0.6, and 1.0 log unit. Bottom row was with the green H filter at intensities of 0.0 and 1.0 log unit below maximum.

Fig. 12
Fig. 12

Sample responses to the FR stimulus of highest intensity. These are filtered responses except in the case of observer JA where the direct-coupled tracing is also shown for comparison.

Fig. 13
Fig. 13

Sample responses to the H stimulus of highest available intensity. With the exception of the bottom row, these are direct-coupled recordings. In the bottom row responses were obtained with the intensity of the test flash reduced 1 log unit.