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Exploring Color Vision: A Comparative Study of Normal and Anomalous Subjects, Study notes of Physiology

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.

What you will learn

  • How does the presence of a weak green light affect the electrical threshold in normal subjects?
  • What are the differences in excitability curves between normal and anomalous subjects?
  • What is the significance of crest times in understanding color vision?

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The Tokoku Journal of Experimental Medicine, Vol . 51, Nos. 3 & 4, 1949.
Physiological Evidence for the Three Components
Theory of Color Vision.
By
Koiti Motokawa.
(本川 一)
(From the Physiological Laboratory of Prof. K. Motokawa,
Tohoku University, Sendai.)
(Received for publication, November 7, 1947)
The theory of Young-Helmholtz is based on the facts of color mixtures.
Young supposed that in the retina there are three types of nerve fibers
which produce a sensation of red, green and blue respectively. The
various color sensations result from the relative strengths with which the
three kinds of fibers are stimulated, and a colorless sensation of white results
if they are all excited equally. Expressed in modern terms, the three
species of nerve fibers assumed by Young may be considered as three
species of cones. Each type of cone contains its own particular photo
sensitive substance, and is concerned with an optic nerve fiber, stimulation
of which produces the color sensation corresponding to it.
Helmholtz1) assumed in each cone three different activities (chemical,
electrical or other processes), because he became aware that the facts of
visual acuity necessitated a smaller unit than three cones. In spite of the
further elaboration of this theory since Helmholtz, there is no physiological
evidence of the existence of three kinds of nerve fibers, cones or activities,
except the experiments of color mixing. The data of color mixing them
selves can be adequately described on the basis of tetrachromatic theories
of Hering2) and others as well, as pointed out by Schrodinger.3) Further
more, these theories are even more natural and simpler from a psychological
point of view; they treat yellow and white as unitary sensations, because
we can perceive no trace of red and green in yellow, and no trace of red,
green and blue in white, although the trichromatic theory claims that
yellow and white are compound sensations.
However, the unitary nature of white as a sensation does not hinder
it from being the result of psychical fusion of two or more separate phy
siological processes. Therefore, it is neither psychological experiments nor
measurements of color mixing that decide the question as to which of these
views is correct. The problem is essentially a physiological one.
The author has shown in a series of papers4)5)6) that the effect of
207
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The Tokoku Journal of Experimental Medicine, Vol. 51, Nos. 3 & 4, 1949.

Physiological Evidence for the Three Components

Theory of Color Vision.

By

Koiti Motokawa.

(From the Physiological Laboratory of Prof. K. Motokawa,

Tohoku University, Sendai.)

(Receivedforpublication,November7, 1947)

The theory of Young-Helmholtz is based on the facts of color mixtures.

Young supposed that in the retina there are three types of nerve fibers

which produce a sensation of red, green and blue respectively. The

various color sensations result from the relative strengths with which the

three kinds of fibers are stimulated, and a colorless sensation of white results

if they are all excited equally. Expressed in modern terms, the three

species of nerve fibers assumed by Young may be considered as three

species of cones. Each type of cone contains its own particular photo

sensitive substance, and is concerned with an optic nerve fiber, stimulation

of which produces the color sensation corresponding to it.

Helmholtz1) assumed in each cone three different activities (chemical,

electrical or other processes), because he became aware that the facts of

visual acuity necessitated a smaller unit than three cones. In spite of the

further elaboration of this theory since Helmholtz, there is no physiological

evidence of the existence of three kinds of nerve fibers, cones or activities,

except the experiments of color mixing. The data of color mixing them

selves can be adequately described on the basis of tetrachromatic theories

of Hering2) and others as well, as pointed out by Schrodinger.3) Further

more, these theories are even more natural and simpler from a psychological

point of view; they treat yellow and white as unitary sensations, because

we can perceive no trace of red and green in yellow, and no trace of red,

green and blue in white, although the trichromatic theory claims that

yellow and white are compound sensations.

However, the unitary nature of white as a sensation does not hinder

it from being the result of psychical fusion of two or more separate phy

siological processes. Therefore, it is neither psychological experiments nor

measurements of color mixing that decide the question as to which of these

views is correct. The problem is essentially a physiological one.

The author has shown in a series of papers4)5)6)that the effect of

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.

  1. Observations on Subjects of Anomalous Color Vision. The general method is described in my preceding papers cited above, so that only points special for this experiment will be mentioned. As shown in my previous work,4) the electrical excitability of the eye as measured by a rheobasic current becomes supernormal after an illumination. The maximum of excitability is reached usually in about two seconds after the end of illumination when white light is used.

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.

if this subject were dichromatic in vision at this intensity. This fact cor

responds to the results of psychological experiments that hue discrimina

tion depends greatly upon the intensities of illumination, and that anomal

ous trichromats become practically dichromatic at low intensities.

Another fact worth while mentioning is that in this subject two dif

ferent values of ordinates were often obtained for one and the same abscissa.

One of the two values fell on one curve, while the other fell on the other

kind of curve. This fact indicates that each curve behaves independently

of the others, and that the fusion of the component curves found in normal

subjects is only apparent one.

2. Demonstration of Three Kinds of Processes

in the Retina of Normal Trichromats.

As seen from Fig. 3 (curves of empty circles and empty triangles), the

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.

  1. 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.

  2. 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

214 K. Motokawa

maxima, one of which has the same crest time as the curve for red light,

and the other of which corresponds to the maximum for blue light.

3. When electrical threshold is determined in the presence of a weak

green light after an illumination with white light, the three maxima appear

in normal subjects like in deuteranomalous persons. This is due to an in

hibitory effect of the green light upon one of the three component-processes.

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.