Professor Hering, of Leipsic, adopting the generally accepted view that light effects chemical changes in substances contained in the retina, to which changes stimulation of nerve-endings is due, formulated a theory of colour-vision which many physiologists prefer to Young’s. He imagines that the retina contains three kinds of pigment, each of which is, as he believes all living substance to be, in a constant state of change. It is at the same time being built up and destroyed. Using the terms which connote the opposite directions of metabolism, the pigment is simultaneously undergoing anabolism and katabolism; the two processes, when the retina is at rest, maintaining equilibrium. When light acts upon either of the substances, it hastens, according to its quality, either the one process or the other; and the chemical change, whether it be constructive or destructive, stimulates the endings of optic nerves. Hering assumes, therefore, that there are six elementary qualities of visual sensation—red, green, yellow, blue, white, black. Red, yellow, white are due to anabolism of the visual substances; green, blue, black are due to their katabolism. The installation of yellow amongst the unanalysable colours is a relief to many minds. It is almost impossible to think of yellow as a compounded colour. White also, we feel, is not a compounded colour, despite our knowledge that a prism scatters from it all the hues of the rainbow. Black, many persons assert, gives them a definite sensation, and not merely a sense of rest. (Parenthetically, it may be observed that the feeling that a colour is pure or mixed is not to be trusted. It may be based upon the chromatic aberration of the eye, or it may be reminiscent of the paint-box. We know that we cannot make yellow by mixing red and green pigments, hence we feel that it is pure. Of green we are not by any means sure; gamboge and Prussian blue come into our minds.) Except when the light which falls upon the retina is giving rise to one of the four pure colour-sensations, all three substances are simultaneously affected, although one may be undergoing katabolism while the other two are being built up, or vice versa. Hering accounts for simultaneous contrast by assuming that the activity of any one part of the retina induces an opposite kind of change in the remainder, and especially in the vicinity of the primarily active part. When a certain patch is developing a sensation of red, the rest of the retina develops a sensation of green.
The great merit of the theory is, however, to be found in its offering an explanation of complementary after-images. The green patch seen with closed eyes after one has stared at a red object is due to the rebound of metabolism. In returning to a condition of chemical equilibrium the retinal substance acts as a stimulant which evokes the antagonistic colour. But it is a theory which makes very large assumptions. It assumes, for example, the possibility of the existence of a substance which is built up by light from one end of the spectrum, and decomposed by light from its centre. Not that Hering regards the existence of three retinal substances as essential to his theory. He is prepared to transfer to the brain the seat of the substances, or the substance, which, by their, or its, anabolism and katabolism, produces antagonistic colour-perceptions; but in this he is abandoning physiology for metaphysics. We have no warrant for imagining that there exists in the brain any substance which, by undergoing physical changes of various kinds, produces various psychical effects. The problem to be solved is physiological. Rays of light of different wave-lengths excite the retina to discharge impulses which are variously distributed in the brain. The effects which they produce in consciousness depend upon their distribution. The impulses to which the longest rays give rise evoke sensations of red, those due to the shortest, sensations of violet. And what is true of the retina as a whole is true, apparently, of each individual cone. In what way does light act upon a cone? It is one of the most fascinating problems in physiology. Round it our thoughts revolve whenever we are trying to form conceptions of the nature of stimulation, sensation, and perception. Each of the two theories which we have expounded above helps to group together certain of the more striking phenomena of colour-vision, but neither gives a satisfying explanation of their causation.
The sensitiveness of the retina is in a remarkable degree adjusted to the intensity of the light. When a dark room is entered, the pupil dilates; but one’s power of distinguishing objects continues to increase after the pupil has reached its maximum size. At the end of ten minutes the eye may be twenty-five times as sensitive as it was when the room was entered. This adaptation to darkness is due in large degree to the substitution of rods for cones as the organs on which vision chiefly depends. But it cannot be wholly due to this, since it occurs when one is working with a red light. Probably the red used in a “dark-room” is not sufficiently near the end of the spectrum to be completely without influence upon visual purple, but it is a colour to which rods are comparatively insensitive. Other evidence also points to an adaptation of cones as well as of rods.
Fig. 32.—The Formation of an Image by the Refracting Media of the Eye.
x, The common centre of curvature (nodal point of the several media). Rays which pass through this point are not deflected. y, The principal focus of the system. All rays which are parallel to the optic axis converge to this point. The image of the point A is formed at a, the spot at which a ray parallel with the optic axis meets an unbent ray—the image of B at b.
Accommodation of the eye for distance is brought about by a mechanism which allows the lens to change in shape. It becomes more convex when a near object is looked at than it was when adjusted for an unlimited distance, which is its condition when the eye is at rest. Adjustment for near objects involves muscular action, and is accompanied by a sense of effort, however slight. Whilst the eye is at rest the lens is mechanically compressed against the anterior layer of its suspensory ligament. Accommodation for near vision is effected by the ciliary muscle, which is placed in the shelf of tissue which projects into the interior of the eyeball. This muscle is made up of a ring of circular fibres, and to the outer side of this, of fibres which radiate backwards and outwards. The longitudinal, or radiating, fibres obtain their purchase by attachment to the firm wall of the globe just beyond the cornea. They spread into the front of the loose chorioid membrane which lines the eye behind the retina. By the joint action of these two sets of plain muscle-fibres the suspensory ligament is slackened, and the extremely elastic lens, previously compressed, bulges forwards. The radius of curvature of its anterior surface changes from 10·3 millimetres for distance to 6 millimetres for vision at the “near point.” It was stated, in connection with the development of the lens ([p. 374]), that the cells of the posterior half of the hollow sphere out of which it is formed grow forwards into extremely long fibres, which traverse its whole thickness. These fibres are bent like the segments of a carriage-spring. Their anterior ends rest against the flattened ligament of the lens; the vitreous humour, which is always under tension, compresses their posterior ends. When removed from the eye, the lens becomes rounder than it is in situ, even when accommodated for near objects. But in later life it grows stiff. It ceases to bulge forwards when its ligament is slackened. Hence it becomes necessary to aid the presbyopic eye with convex glasses when it is used for near objects, although for distant vision it remains as effective as ever. If the ciliary muscle is constantly and completely relieved of the labour of accommodation, it grows lazy, or rather wastes from want of use. A person who relies on spectacles loses his power of accommodation; but ophthalmologists agree that self-focussing, if it give rise to a sensation of strain, is bad for the eyes. In myopic persons the eyeball is too deep; objects are focussed in front of the retina. In hypermetropia (“long sight”) the eyeball is too shallow; objects are focussed behind the retina. Concave glasses correct the one condition, convex glasses correct the other. Glasses are also very commonly called for to neutralize another defect—regular astigmatism—which may be present by itself, or may accompany insufficient length or too great length of the optic axis. It is due to unequal curvature of the cornea. Usually the curvature is sharper in the vertical than in the horizontal meridian ([cf. p. 269]); as a consequence, points in a vertical line are focussed in front of points in a horizontal line. Cylindrical glasses, not lenses, are required to correct this defect. And here it may be well to call attention to the fact that rays of light are more sharply refracted by the surface of the cornea than they are by the crystalline lens. The lens has a high index of refraction (1·45), but it does not lie in air (the index of refraction of which is 1), but between two humours which have about the same index as water—namely, 1·336. The bending by the combined action of the cornea and the lens of rays of light which come from a source so distant that they may be considered as parallel brings them to a focus on the retina, when the lens is at its flattest. When the lens is at its roundest, rays which diverge from a point only 5 inches in front of the eye are focussed on the retina. The lens is therefore essential for accommodation, but, after its removal for cataract, vision, even for near objects, is rendered possible by the use of convex glasses.