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Insights into the physiological evidence for the trichromatic theory of color vision. It discusses the differences in excitability curves between normal and anomalous subjects, specifically those with deuteranomalous vision. The document also explains the concept of crest times and their significance in understanding color vision.
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The Tokoku Journal of Experimental Medicine, Vol. 51, Nos. 3 & 4, 1949.
208 K. Motokawa
colors upon the retina shows itself in changes of the electrical excitability of the retina. In the present communication, this phenomenon has been made use of to prove that in reality, there are three kinds of physiological processes in the retina, as the three components-theory assumes.
Results.
Fig. 1, Excitability curve of the eye of a deuteranomalous person (K. A.) for white light. Ordinates: Increases in electrical excitability (reciprocal of threshold) in percentage of the excitability in the dark without preceding illumination. Abscissae: Time intervals between the end of illumination two seconds in duration and the stimula tion with a constant current 100 msec. in duration. It has been found that the time course of electrical excitability is quite different in persons of anomalous color vision from that in normal subjects. Particularly interesting for the present problem are the curves obtained from deuteranomalous persons, for they have three maxima, as if they corresponded to the three component-processes as hypothesized in the trichromatic theory. An example is illustrated in Fig. 1. The illumina tion was of an intensity 10 lux and confined to the center of the eye (the visual angle 2•‹ in diameter). If the time interval between the end of illumination and the maximum of excitability is designated "crest time,"
210 K. Motokawa
Fig. 2. The effect of intensities of white light upon the excitability curves at a deuteranomalous subject (K. S.) Ordinates: Excitability increases due to illumination with white light in percentage of the excitability without preceding illumination. Abscissa: Intervals between the end of illumination and electrical stimulation.
PhysiologicalEvidencefor TrichromaticTheory 211
excitability curves for white light have only a single maximum ause the three processes R , probably bec , G and B are excited in such proportions as the three curves apparently fuse together into a congruent curve. If such proportions are unbalanced in one way or another, the fusion will become less complete, and consequently the three maxima will be disclosed.
Fig. 3. Experiments showing the existence of three kinds of retinal processes in normal trichromats. (A): Excitability curve for white light (empty circles) and that for a monochromatic light 510mƒÊ in wave-length (solid circles). Subject: T. M. ( B): Usual excitability curve for white light (empty triangles) and the curve for white light at one and the same subject under the influence of a weak green light (530mƒÊ) at the moment of electrical stimulation. The G-component is partially inhibited by the green light, and consequently the three maxima appear. Double circles and double triangles indicate the coincidence of two measurements.
According to the three components-theory, the relative strengths with which the three components are excited, must be different from wave length to wave-length. Therefore, the three maxima must be revealed by illuminating with a monochromatic light of some adequate wave-length.
PhysiologicalEvidencefor TrichromaticTheory 213
The curve shows distinct three maxima, the crest times of which are 1, 2.25 and 3 seconds. These values coincide with those of the previous case,
suggesting that there is little individual difference among normal tri chromats. There is now no doubt that the appearance of three maxima in deuter anomalous persons is due to the congenital weakness of the G-component, for it has experimentally proved that three maxima are disclosed by partial inhibition of the G-component in normal trichromats in whom otherwise only a monophasic curve is obtained for white light. In subjects of anomalous color vision, not only the proportions of the three processes are less harmonious, but also the values of crest times de viate greatly from those of normal trichromats. At any rate, our findings are in entire harmony with the theory of Young and Helmholtz, whereas they are contradictory to the theory of Hering at least in regard to the preripheral mechanism; our findings show that the process for white can be analysed into three component-processes against Hering's view that there should be a special substance in the retina for the reception of white light. However, such discussion should be restricted to the findings at the fovea, for data obtained at the periphery seem to agree neither with the theory of Young-Helmholtz nor with that of Hering, as will be reported in detail elsewhere. SUMMARY.
The electrical excitability of the human eye was measured at varying intervals after the end of an illumination two seconds in duration by means of a rheobasic constant current, and the differences between the excitabilities with and without the preceding illumination were plotted as ordinates against time intervals between the end of illumination and electrical stimula tion as abscissas. The illumination was restricted to the fovea centralis.
The curve so obtained has a single maximum in normal trichromats, but three maxima in deuteranomalous subjects when white light is used for illumination. The height of each maximum depends greatly upon intensities of illumination, but the crest time is entirely independent of intensities. The appearance of the three maxima is so interpreted that one of the three component-processes is too weak for such complete fusion as is found at normal subjects.
In some normal trichromats the three maxima appear in the ex citability curve for a monochromatic light of some adequate wave-length. It was found that bluish green light is adequate for this purpose. Red light gives rise to a excitability curve with a single maximum, the crest time of which is about 1 second. The curve for blue light is also mono phasic, the crest time being about 3 seconds. Violet light causes two
All above findings provide physiological evidence of the three com ponents-theory of Young-Helmholtz.
References. (1) Helmholtz, H., Handbuch der physiologischen Optik (3 rd ed.) 3 Vols,. Hamburg & Leipzig. (2) Hering, E., Grundzuge der Lehre vom Lichtsinn. Berlin. (3) Schrodinger, E., Sitzgsber. d. Wien. Akad. d. Wiss., Math.•\Nat. Kl., 1925, 134, ii a, 471. (4) Motokawa, K., Tohoku J. Exp. Med., 1949, 51, 165. (5) Motokawa, K., Tohoku J. Exp. Med., 1949, 51, 179. (6) Motokawa, K., Tohoku J. Exp. Med., 1949, 51, 197.