"In the nystagmus," he continues, "the eyeballs execute two well-differentiated motions: one, a compensatory, relatively slow motion, during which images pass across the retina so as to give the appearance of a movement of space in the opposite direction; two, a swift motion opposite to the slow one, and so rapid that the images passing across the retina leave no sensation of their movement."
Now, in a previous paper[17] I have shown that there is a central anæsthesia, or central inhibition of visual sensations, during about the latter two thirds of the time occupied by every voluntary eye-jump; and in view of this I was led to enquire whether in fact, as Delage so confidently asserts, it is the speed of these more rapid movements, or some other factor, that causes them to leave no visual sensations. There can be no doubt that they do leave none, since, aside from the statement of Delage, in dizziness the visual field whirls always in only one direction; whereas it should otherwise appear to swing now to one side, now to the other, as the eyes move back and forth across the objects. I have found but one other mention of this point in the literature. In his Analyse,[18] Mach says, parenthetically, "(the jerky eye-movement leaves no optical impression)"; but he does not suggest that this is because of its greater speed.
In order to test this point, a 2 c. p. incandescent lamp was so arranged that it could be moved vertically in front of, and about four metres distant from, a rotating chair. Since after a rotation the eyes are oscillating from side to side, if the lamp is moved up and down an obliquely inclined after-image streak must be generated on the retina; and clearly there are four possible positions in which this may lie, as shown in Fig. 1.
The results were absolutely uniform (the author alone as subject); the after-image streak always lay on that side of the moving light toward which the slow eye-movements were directed, that is, the lamp appeared to drift obliquely up or down and in a lateral direction opposite to that of the slow eye-movements. Apart from its vertical displacement, then, the lamp behaved like the less intensely illumined parts of the visual field, seeming to be totally invisible during the swifter eye-movements. Now since the experiment was done in a partially darkened room and the eyes were partly adapted to darkness, the lamp should have been intense enough adequately to stimulate the retina even during the more rapid movements, and might be expected to leave an after-image streak on that side toward which these rapid movements were directed, and differing only from the streaks seen during the slow eye-movements in being inclined at a less angle from the horizontal. Yet no such streaks were visible.
These observations were made at about the same number of seconds after the rotation stopped, as the photographs were taken that are recorded in the preceding paper of this volume. The rapid movements were therefore about one sixth as long in duration as the slower ones. Since the respective amplitudes of rapid and slow must average very nearly the same, the rapid movements must have been about six times as swift as the slow movements. It needs therefore to be shown beyond a doubt that the 2 c. p. lamp was bright enough, in view of the briefness of stimulation of any one retinal element during the rapid eye-movement, to be above the threshold of perception. For this reason the experiment was not continued with other subjects.
The certainly adequate degree of illumination was realized during the photography of the eyes described in the preceding paper. Here during the post-rotary dizziness an arc lamp (of 6 amp.) was in front of the face and but a little to one side of the primary line of regard; it was 60 cm. distant from the eyes and on a level with them; a lens condensed the rays on the two eyes, and the light was diminished only just enough as not to be painful, by a dilute screen of copper ammonium sulphate about 3 cm. thick. Of course such an illumination must adequately stimulate each retinal element even during the most rapid eye-movements. Nevertheless with the four subjects that were photographed the arc lamp, like the rest of the visual field, seemed always to swim in one direction, and that opposite to the slower eye-movements. In one case where the eyes were photographed without the adiathermal screen, and the light was rather painfully intense, the lamp was still seen to drift in one and the same direction. There was never any trace of its moving to and fro, as there should have been had it been visible during both phases of the nystagmiform movements.
Fig. 1
This absence of visual sensation during the more rapid eye-movements might conceivably depend on either peripheral or central inhibitory factors. But the anatomy and physiology of the eye offer no point of support for the supposition that during such movements the irritability of the rods and cones is momentarily reduced, or that the retinal layers posterior to the rods and cones suffer an interruption of function during a movement of the eyeball in its socket. Indeed, during some such movements, the "pursuit" movements (Dodge's second type), vision is unimpaired.[19] In view of these facts, and of the many known cases of the mutual inhibition of sensations where undoubtedly the process is a central one, it is by far most probable that this visual inhibition is also a central process; as was certainly the visual inhibition during voluntary eye-jumps, previously reported by me.[20]