Sight.

Ferrier was the first in the field here. He found, when the angular convolution (that lying between the 'intra parietal' and 'external occipital' fissures, and bending round the top of the fissure of Sylvius, in Fig. 6) was excited in the monkey, that movements of the eyes and head as if for vision occurred; and that when it was extirpated, what he supposed to be total and permanent blindness of the opposite eye followed. Munk almost immediately declared total and permanent blindness to follow from destruction of the occipital lobe in monkeys as well as dogs, and said that the angular gyrus had nothing to do with sight, but was only the centre for tactile sensibility of the eyeball. Hunk's absolute tone about his observations and his theoretic arrogance have led to his ruin as an authority. But he did two things of permanent value. He was the first to distinguish in these vivisections between sensorial and psychic blindness, and to describe the phenomenon of restitution of the visual function after its first impairment by an operation; and the first to notice the hemiopic character of the visual disturbances which result when only one hemisphere is injured. Sensorial blindness is absolute insensibility to light; psychic blindness is inability to recognize the meaning of the optical impressions, as when we see a page of Chinese print but it suggests nothing to us. A hemiopic disturbance of vision is one in which neither retina is affected in its totality, but in which, for example, the left portion of each retina is blind, so that the animal sees nothing situated in space towards its right. Later observations have corroborated this hemiopic character of all the disturbances of sight from injury to a single hemisphere in the higher animals; and the question whether an animal's apparent blindness is sensorial or only psychic has, since Munk's first publications, been the most urgent one to answer, in all observations relative to the function of sight.

Goltz almost simultaneously with Ferrier and Munk reported experiments which led him to deny that the visual function was essentially bound up with any one localized portion of the hemispheres. Other divergent results soon came in from many quarters, so that, without going into the history of the matter any more, I may report the existing state of the case as follows:[17]

In fishes, frogs, and lizards vision persists when the hemispheres are entirely removed. This is admitted for frogs and fishes even by Munk, who denies it for birds.

All of Munk's birds seemed totally blind (blind sensorially) after removal of the hemispheres by his operation. The following of a candle by the head and winking at a threatened blow, which are ordinarily held to prove the retention of crude optical sensations by the lower centres in supposed hemisphereless pigeons, are by Munk ascribed to vestiges of the visual sphere of the cortex left behind by the imperfection of the operation. But Schrader, who operated after Munk and with every apparent guarantee of completeness, found that all his pigeons saw after two or three weeks had elapsed, and the inhibitions resulting from the wound had passed away. They invariably avoided even the slightest obstacles, flew very regularly towards certain perches, etc., differing toto cœlo in these respects with certain simply blinded pigeons who were kept with them for comparison. They did not pick up food strewn on the ground, however. Schrader found that they would do this if even a small part of the frontal region of the hemispheres was left, and ascribes their non-self-feeding when deprived of their occipital cerebrum not to a visual, but to a motor, defect, a sort of alimentary aphasia.[18]

In presence of such discord as that between Munk and his opponents one must carefully note how differently significant is loss, from preservation, of a function after an operation on the brain. The loss of the function does not necessarily show that it is dependent on the part cut out; but its preservation does show that it is not dependent: and this is true though the loss should be observed ninety-nine times and the preservation only once in a hundred similar excisions. That birds and mammals can be blinded by cortical ablation is undoubted; the only question is, must they be so? Only then can the cortex be certainly called the 'seat of sight.' The blindness may always be due to one of those remote effects of the wound on distant parts, inhibitions, extensions of inflammation,—interferences, in a word,—upon which Brown-Séquard and Goltz have rightly insisted, and the importance of which becomes more manifest every day. Such effects are transient; whereas the symptoms of deprivation (Ausfallserscheinungen, as Goltz calls them) which come from the actual loss of the cut-out region must from the nature of the case be permanent. Blindness in the pigeons, so far as it passes away, cannot possibly be charged to their seat of vision being lost, but only to some influence which temporarily depresses the activity of that seat. The same is true mutatis mutandis of all the other effects of operations, and as we pass to mammals we shall see still more the importance of the remark.

In rabbits loss of the entire cortex seems compatible with the preservation of enough sight to guide the poor animals' movements, and enable them to avoid obstacles. Christiani's observations and discussions seem conclusively to have established this, although Munk found that all his animals were made totally blind.[19]

In dogs also Munk found absolute stone-blindness after ablation of the occipital lobes. He went farther and mapped out determinate portions of the cortex thereupon, which he considered correlated with definite segments of the two retinæ, so that destruction of given portions of the cortex produces blindness of the retinal centre, top, bottom, or right or left side, of the same or opposite eye. There seems little doubt that this definite correlation is mythological. Other observers, Hitzig, Goltz, Luciani, Loeb, Exner, etc., find, whatever part of the cortex may be ablated on one side, that there usually results a hemiopic disturbance of both eyes, slight and transient when the anterior lobes are the parts attacked, grave when an occipital lobe is the seat of injury, and lasting in proportion to the latter's extent. According to Loeb, the defect is a dimness of vision ('hemiamblyopia') in which (however severe) the centres remain the best seeing portions of the retina, just as they are in normal dogs. The lateral or temporal part of each retina seems to be in exclusive connection with the cortex of its own side. The centre and nasal part of each seems, on the contrary, to be connected with the cortex of the opposite hemispheres. Loeb, who takes broader views than any one, conceives the hemiamblyopia as he conceives the motor disturbances, namely, as the expression of an increased inertia in the whole optical machinery, of which the result is to make the animal respond with greater effort to impressions coming from the half of space opposed to the side of the lesion. If a dog has right hemiamblyopia, say, and two pieces of meat are hung before him at once, he invariably turns first to the one on his left. But if the lesion be a slight one, shaking slightly the piece of meat on his right (this makes of it a stronger stimulus) makes him seize upon it first. If only one piece of meat be offered, he takes it, on whichever side it be.

When both occipital lobes are extensively destroyed total blindness may result. Munk maps out his 'Sehsphäre' definitely, and says that blindness must result when the entire shaded part, marked A, A, in Figs. 12 and 13, is involved in the lesion. Discrepant reports of other observations he explains as due to incomplete ablation. Luciani, Goltz, and Lannegrace, however, contend that they have made complete bilateral extirpations of Munk's Sehsphäre more than once, and found a sort of crude indiscriminating sight of objects to return in a few weeks.[20] The question whether a dog is blind or not is harder to solve than would at first appear; for simply blinded dogs, in places to which they are accustomed, show little of their loss and avoid all obstacles; whilst dogs whose occipital lobes are gone may run against things frequently and yet see notwithstanding. The best proof that they may see is that which Goltz's dogs furnished: they carefully avoided, as it seemed, strips of sunshine or paper on the floor, as if they were solid obstacles. This no really blind dog would do. Luciani tested his dogs when hungry (a condition which sharpens their attention) by strewing pieces of meat and pieces of cork before them. If they went straight at them, they saw; and if they chose the meat and left the cork, they saw discriminatingly. The quarrel is very acrimonious; indeed the subject of localization of functions in the brain seems to have a peculiar effect on the temper of those who cultivate it experimentally. The amount of preserved vision which Goltz and Luciani report seems hardly to be worth considering, on the one hand; and on the other, Munk admits in his penultimate paper that out of 85 dogs he only 'succeeded' 4 times in his operation of producing complete blindness by complete extirpation of his 'Sehsphäre.'[21] The safe conclusion for us is that Luciani's diagram, Fig. 14, represents something like the truth. The occipital lobes are far more important for vision than any other part of the cortex, so that their complete destruction makes the animal almost blind. As for the crude sensibility to light which may then remain, nothing exact is known either about its nature or its seat.

In the monkey, doctors also disagree. The truth seems, however, to be that the occipital lobes in this animal also are the part connected most intimately with the visual function. The function would seem to go on when very small portions of them are left, for Ferrier found no 'appreciable impairment' of it after almost complete destruction of them on both sides. On the other hand, he found complete and permanent blindness to ensue when they and the angular gyri in addition were destroyed on both sides. Munk, as well as Brown and Schaefer, found no disturbance of sight from destroying the angular gyri alone, although Ferrier found blindness to ensue. This blindness was probably due to inhibitions exerted in distans, or to cutting of the white optical fibres passing under the angular gyri on their way to the occipital lobes. Brown and Schaefer got complete and permanent blindness in one monkey from total destruction of both occipital lobes. Luciani and Seppili, performing this operation on two monkeys, found that the animals were only mentally, not sensorially, blind. After some weeks they saw their food, but could not distinguish by sight between figs and pieces of cork. Luciani and Seppili seem, however, not to have extirpated the entire lobes. When one lobe only is injured the affection of sight is hemiopic in monkeys: in this all observers agree. On the whole, then, Munk's original location of vision in the occipital lobes is confirmed by the later evidence.[22]

In man we have more exact results, since we are not driven to interpret the vision from the outward conduct. On the other hand, however, we cannot vivisect, but must wait for pathological lesions to turn up. The pathologists who have discussed these (the literature is tedious ad libitum) conclude that the occipital lobes are the indispensable part for vision in man. Hemiopic disturbance in both eyes comes from lesion of either one of them, and total blindness, sensorial as well as psychic, from destruction of both.

Hemiopia may also result from lesion in other parts, especially the neighboring angular and supra-marginal gyri, and it may accompany extensive injury in the motor region of the cortex. In these cases it seems probable that it is due to an actio in distans, probably to the interruption of fibres proceeding from the occipital lobe. There seem to be a few cases on record where there was injury to the occipital lobes without visual defect. Ferrier has collected as many as possible to prove his localization in the angular gyrus.[23] A strict application of logical principles would make one of these cases outweigh one hundred contrary ones. And yet, remembering how imperfect observations may be, and how individual brains may vary, it would certainly be rash for their sake to throw away the enormous amount of positive evidence for the occipital lobes. Individual variability is always a possible explanation of an anomalous case. There is no more prominent anatomical fact than that of the 'decussation of the pyramids,' nor any more usual pathological fact than its consequence, that left-handed hemorrhages into the motor region produce right-handed paralyses. And yet the decussation is variable in amount, and seems sometimes to be absent altogether.[24] If, in such a case as this last, the left brain were to become the seat of apoplexy, the left and not the right half of the body would be the one to suffer paralysis.

The schema below [Fig. 15], copied from Dr. Seguin, expresses, on the whole, the probable truth about the regions concerned in vision. Not the entire occipital lobes, but the so-called cunei, and the first convolutions, are the cortical parts most intimately concerned. Nothnagel agrees with Seguin in this limitation of the essential tracts.[25]

A most interesting effect of cortical disorder is mental blindness. This consists not so much in insensibility to optical impressions, as in inability to understand them. Psychologically it is interpretable as loss of associations between optical sensations and what they signify; and any interruption of the paths between the optic centres and the centres for other ideas ought to bring it about. Thus, printed letters of the alphabet, or words, signify certain sounds and certain articulatory movements. If the connection between the articulating or auditory centres, on the one hand, and the visual centres on the other, be ruptured we ought a priori to expect that the sight of words would fail to awaken the idea of their sound, or the movement for pronouncing them. We ought, in short, to have alexia, or inability to read: and this is just what we do have in many cases of extensive injury about the fronto-temporal regions, as a complication of aphasic disease. Nothnagel suggests that whilst the cuneus is the seat of optical sensations, the other parts of the occipital lobe may be the field of optical memories and ideas, from the loss of which mental blindness should ensue. In fact, all the medical authors speak of mental blindness as if it must consist in the loss of visual images from the memory. It seems to me, however, that this is a psychological misapprehension. A man whose power of visual imagination has decayed (no unusual phenomenon in its lighter grades) is not mentally blind in the least, for he recognizes perfectly all that he sees. On the other hand, he may be mentally blind, with his optical imagination well preserved; as in the interesting case published by Wilbrand in 1887.[26] In the still more interesting case of mental blindness recently published by Lissauer,[27] though the patient made the most ludicrous mistakes, calling for instance a clothes-brush a pair of spectacles, an umbrella a plant with flowers, an apple a portrait of a lady, etc. etc., he seemed, according to the reporter, to have his mental images fairly well preserved. It is in fact the momentary loss of our non-optical images which makes us mentally blind, just as it is that of our non-auditory images which makes us mentally deaf. I am mentally deaf if, hearing a bell, I can't recall how it looks; and mentally blind if, seeing it, I can't recall its sound or its name. As a matter of fact, I should have to be not merely mentally blind, but stone-blind, if all my visual images were lost. For although I am blind to the right half of the field of view if my left occipital region is injured, and to the left half if my right region is injured, such hemianopsia does not deprive me of visual images, experience seeming to show that the unaffected hemisphere is always sufficient for production of these. To abolish them entirely I should have to be deprived of both occipital lobes, and that would deprive me not only of my inward images of sight, but of my sight altogether.[28] Recent pathological annals seem to offer a few such cases.[29] Meanwhile there are a number of cases of mental blindness, especially for written language, coupled with hemianopsia, usually of the rightward field of view. These are all explicable by the breaking down, through disease, of the connecting tracts between the occipital lobes and other parts of the brain, especially those which go to the centres for speech in the frontal and temporal regions of the left hemisphere. They are to be classed among disturbances of conduction or of association; and nowhere can I find any fact which should force us to believe that optical images need[30] be lost in mental blindness, or that the cerebral centres for such images are locally distinct from those for direct sensations from the eyes.[31]

Where an object fails to be recognized by sight, it often happens that the patient will recognize and name it as soon as he touches it with his hand. This shows in an interesting way how numerous the associative paths are which all end by running out of the brain through the channel of speech. The hand-path is open, though the eye-path be closed. When mental blindness is most complete, neither sight, touch, nor sound avails to steer the patient, and a sort of dementia which has been called asymbolia or apraxia is the result. The commonest articles are not understood. The patient will put his breeches on one shoulder and his hat upon the other, will bite into the soap and lay his shoes on the table, or take his food into his hand and throw it down again, not knowing what to do with it, etc. Such disorder can only come from extensive brain-injury.[32]

The method of degeneration corroborates the other evidence localizing the tracts of vision. In young animals one gets secondary degeneration of the occipital regions from destroying an eyeball, and, vice versâ, degeneration of the optic nerves from destroying the occipital regions. The corpora geniculata, thalami, and subcortical fibres leading to the occipital lobes are also found atrophied in these cases. The phenomena are not uniform, but are indisputable;[33] so that, taking all lines of evidence together, the special connection of vision with the occipital lobes is perfectly made out. It should be added, that the occipital lobes have frequently been found shrunken in cases of inveterate blindness in man.