Colour vision

by Sir William de Wiveleslie Abney

Scl is the sclerotic coat. Ch the choroid coat, with CP the ciliary process. I is the body of the Iris. R is the retina or inner wall. PE the pigment epithelium or outer wall. L the lens held by the suspensory ligament sp.l. VH is the vitreous humour. ON the optic nerve, ox is the optic axis, in this case made to pass through the fovea centralis, f.c.

As far as the perception of colour is concerned, the principal part of the eye which is not distinctly optical—i.e. for the production of images—is the retina, and this it will be seen is in reality an outcrop of the brain, the connection between the two being the optic nerve. Owing to this connection, it is not easy to determine where the seat of colour perception is located; but for the purpose of physical investigation this is not of first-rate importance, nor does it affect the discussion of rival theories except in a minor degree. There are other subsidiary adjuncts in the eye to which, however, I must call attention, as they have a distinct bearing on the apparent intensity of some colours and of the hue that mixtures of others are perceived. The first is what is called the “macula lutea,” or yellow spot, a spot which it may be assumed exists in every eye. It is horizontally oval in form, and is situated in the very centre of the retina, embracing some 6° to 8° in angular measure. It has a brownish or yellowish tint, and the retina at this part is slightly depressed, being bounded by a slightly raised rim. In the centre of this area the retina becomes very thin, having a depression about 1/100 of an inch or ·3 millimetres in diameter, which is named the “fovea centralis,” where it is said that vision is the most acute. This statement can be well credited when we come to consider where the seat of the stimulation of sensation lies. The colour which tints the yellow spot is strongest at the crater-like rim, and fades away centrally and peripherally, and is said to be wholly absent in the fovea centralis.

As the colour of this spot is yellow or brown in the living eye (and that it is probably brown the absorption indicates), it follows that white light passing through it must be deprived of some of its components, though in differing degrees. If the seat of sensation is at the outer layer of the retina, as we shall shortly see must be the case, it will further be seen that when light of any colour which the brown pigment will absorb more or less completely falls on different parts of the oval area, the absorption must vary at each part, and the intensity of the perceived light will be least at the rim and increase centrally and peripherally. As the centre of the yellow spot or fovea is coincident approximately with the point where the axis of the eye cuts the retina, the image of an evenly illuminated object, when looked at directly, must fall on the yellow spot. If, therefore, a patch of such light, the image of which more than covers the spot, be observed, it ought to exhibit a varying brightness of colour corresponding to the strength of the colouring matter which exists at the different parts. This it but rarely does, for habit and constant interpretation of what should be seen prevents the mind from distinguishing these variations; but if the colour brightness, as perceived by the different parts, be submitted to measurement by proper means, the variations in brightness of the image can be readily recognised. A very common method of exhibiting the presence of the pigment is to look at a bright white cloud through a layer of chrome alum. Chrome alum transmits red and blue-green rays. Now as the spectrum-blue rays are those which the pigment will absorb, it follows that the colour of the solution should appear ruddy to the central part of the eye, but on the rest of the retina it should appear of its ordinary purplish colour. At a first glance, and before the eye has become fatigued, this is the case, but the phenomenon soon disappears. Another way of forming an idea as to what the yellow spot absorbs is to throw a feeble spectrum on a white surface and cause the eye to travel along it. If the spectrum be viewed so that it does not occupy more than about 40° of the retina, the movement of the eye will show a dark band travelling along the green, blue, and violet regions as the image of these parts of the spectrum fall on the yellow spot, and their apparent brightness will increase as they fall outside the absorbing area. This proves that an absorption takes place in this area.

Fig. 2.

The retina consists essentially of an inner and outer wall, enclosing matter which is similar to the grey matter of the brain. On the inner wall are the vessels which are connected with the optic nerve. The outer wall is epithelium coloured with a pigment, and it is here that the visual impulses begin, although the rays of light giving rise to them have to pass through the thickness of the retina before so doing. It has already been stated that the light has to pass through the thickness of the yellow spot before a visual sensation is felt in the centre of the field, and the experiments just given offer a fair proof of the truth of the assertion, but there is still another which is perhaps more conclusive. Suppose we have a hollow reflecting ball, as shown in [Fig. 2], and through an orifice A we project a beam of light to B, which meets an obstruction, S, in its path, then A B would be reflected from B to C on a screen C F, and the obstruction S would be marked at C. If another beam from D was directed so as to meet the same obstruction, its presence would be marked at F. Knowing the distance of the centre O of the hollow sphere from F C and its diameter, and measuring the distance between F and C and their respective distances from the axis of the sphere, the distances S B and S E can be calculated. This method is applied in the formation of what are known as Purkinje’s figures. The simplest case is where a beam of light is directed through the sclerotic and transmitted through the lens. Images of the retinal vessels are distinguished as at S, and it is found that they cast shadows, which are seen as dark lines in the glare of the field of vision. The sensation of light must therefore come from behind these vessels, and calculation shows that the seat of the sensation is close to the pigmented inner wall of the retina.

Lying here is a layer of what are known as rods and cones, which have a connection, either actual or functional, with the optic fibres which largely compose the inner wall of the retina, and are connected with the optic nerve. In the yellow spot the cones are much more numerous than the rods, but in the peripheral part the reverse is the case. In the fovea the rods appear to be altogether absent. The total number of cones in the eye has been calculated to be about 3,000,000, of which about 7,000 are in the small fovea. The number of cones will give an idea of their dimensions. This detail has been entered into as it has been supposed that these rods and cones are all-important in translating light-waves into visual impulses. The inner wall of the retina of most human eyes, as has been mentioned, is stained with a black pigment, fuscin, though in albinos it is absent. What its particular use may be is still unknown, for its change by light is so slow that it can scarcely be the cause of vision. In the outer parts of the rods is, however, diffused a substance highly sensitive to light, called the “visual purple,” from its colour, and a theory founded on chemical action, produced by a change in this substance, has been promulgated. Fascinating, however, as such a theory must be, it lacks confirmation. The fact that the cones do not contain it, and that in the fovea are cones alone, renders it difficult to reconcile the theory with the fact that this part of the retina possesses, we are told, the greatest acuteness of sensation as regards light and colour.

The eyes of most vertebrate animals, it may be remarked, have this visual purple, but in those of the bat, owl, hen, and some others the colouring matter seems to be absent. Visual purple is an interesting substance, however, and as it is found in the eye it probably exercises some useful function, though what that function may be is at present unknown. That images of objects can be formed on the retina, owing to the bleaching of this substance, has been proved by experiment. The purple is first changed to a yellow colour, and then passes into white. These “optograms,” as they are called, can be fixed in an excised eye if the retina be detached, and then be treated with a weak solution of alum.

Fig. 3.