Opposite effects of large and small doses.—There remains a very curious phenomenon, known not only to students of physiological response but also known in medical practice, namely that of the opposite effects produced by the same reagent when given in large or in small doses. Here, too, we have the same phenomena reproduced in an extraordinary manner in inorganic response. The same reagent which becomes a ‘poison’ in large quantities may act as a stimulant when applied in small doses. This is seen in record [fig. 94], in which (a) gives the normal responses in water; KHO solution was now added so as to make the strength three parts in 1,000, and (b) shows the consequent enhancement of response. A further quantity of KHO was added so as to increase the strength to three parts in 100. This caused a complete abolition (c) of response.
It will thus be seen that as in the case of animal tissues and of plants, so also in metals, the electrical responses are exalted by the action of stimulants, lowered by depressants, and completely abolished by certain other reagents. The parallelism will thus be found complete in every detail between the phenomena of response in the organic and the inorganic.
CHAPTER XVII
ON THE STIMULUS OF LIGHT AND RETINAL CURRENTS
- Visual impulse: (1) chemical theory; (2) electrical theory
- —Retinal currents
- —Normal response positive
- —Inorganic response under stimulus of light
- —Typical experiment on the electrical effect induced by light.
The effect of the stimulus of light on the retina is perceived in the brain as a visual sensation. The process by which the ether-wave disturbance causes this visual impulse is still very obscure. Two theories may be advanced in explanation.
(1) Chemical theory.—According to the first, or chemical, theory, it is supposed that certain visual substances in the retina are affected by light, and that vision originates from the metabolic changes produced in these visual substances. It is also supposed that the metabolic changes consist of two phases, the upward, constructive, or anabolic phase, and the downward, destructive, or katabolic phase. Various visual substances by their anabolic or katabolic changes are supposed to produce the variations of sensation of light and colour. This theory, as will be seen, is very complex, and there are certain obstacles in the way of its acceptance. It is, for instance, difficult to see how this very quick visual process could be due to a comparatively slow chemical action, consisting of the destructive breaking-down of the tissue, followed by its renovation. Some support was at first given to this chemical theory by the bleaching action of light on the visual purple present in the retina, but it has been found that the presence or absence of visual purple could not be essential to vision, and that its function, when present, is of only secondary importance. For it is well known that in the most sensitive portion of the human retina, the fovea centralis, the visual purple is wanting; it is also found to be completely absent from the retinæ of many animals possessing keen sight.
(2) Electrical theory.—The second, or electrical, theory supposes that the visual impulse is the concomitant of an electrical impulse; that an electrical current is generated in the retina under the incidence of light, and that this is transmitted to the brain by the optic nerve. There is much to be said in favour of this view, for it is an undoubted fact, that light gives rise to retinal currents, and that, conversely, an electrical current suitably applied causes the sensation of light.
Retinal currents.—Holmgren, Dewar, McKendrick, Kuhne, Steiner, and others have shown that illumination produces electric variation in a freshly excised eye. About this general fact of the electrical response there is a widespread agreement, but there is some difference of opinion as regards the sign of this response immediately on the application, cessation, and during the continuance of light. These slight discrepancies may be partly due to the unsatisfactory nomenclature—as regards use of terms positive and negative—hitherto in vogue and partly also to the differing states of the excised eyes observed.
Waller, in his excellent and detailed work on the retinal currents of the frog, has shown how the sign of response is reversed in the moribund condition of the eye.