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Experiments on special sense localisations in the cortex cerebri of the monkey. Brain 1888: 10; 362380; and Experiments on the electrical excitation of the visual area of the cerebral cortex in the monkey. By E. A. Schäfer FRS, Jodrell Professor of Physiology in University College, London. Brain 1888: 11; 16; with Schäfer on the temporal and occipital lobes. By David Ferrier, MD, FRS. Brain 1888: 11; 730.Localization of cerebral function excited much heated exchange in the 1880s. In 1879, Luciani and Tamburini had analysed 41 clinical cases and concluded that the psycho-sensory centres of vision are situated both in the angular gyrus and in the occipital lobe ... reconciling the rival theories of Ferrier and Munk, and [getting] rid of the[ir] negative observations (Abstracts of British and Foreign Journals. On the sensory centres in the cortex cerebri. Studi clinici sui centri corticali, comunicazione preventive dei Professori Luigi Luciani e Augusto Tamburini, Milan. 1879. W. W. Ireland. Brain 1880: 2; 5801). Two other giants clashed in the April 1888 issue of Brain.
There is no doubt that Edward Albert Schäfer (later Sir Edward Sharpey-Schäfer) started the trouble. In the January 1888 issue of Brain, Schäfer waved a red rag at his opponent: [David] Ferrier ... originally denied the participation of the occipital lobe in the visual perceptive function, and still appears to regard it as subordinate to the angular gyrus. And his own position was clear: removal of both occipital lobes produces total and permanent blindness, whereas destruction of the cortex of both angular gyri is not followed by any appreciable permanent defect of vision. Whereas Schäfer's original work with (Sir) Victor Horsley had indeed indicated that unilateral occipital lesions in the monkey only produce transient crossed hemianopia (confusingly described as bilateral homonymous hemianopsia), these lesions were not complete. Hence, the data could not of themselves absolutely refute the claim by Ferrier that the angular gyrus is the special region of clear or central vision of the opposite eye, and perhaps to some extent also of the eye on the same side. Therefore, with Dr Sanger Brown, Schäfer produced lesions of the angular gyrus or the occipital lobe, unilaterally or on both sides, alone and in combination. Now, separating the lips of the fissures and scooping away the entire gyrus angularis of one side produced results that did not match Ferrier's observations. Much hinged on the observation of eye movements. As for the angular gyrus, Schäfer concluded: loss of vision only occurred when the lesion was so extensive as [also] to disrupt the vascular distribution of the occipital lobe. Nor did the experiments support the view of Professor Munk that the angular gyrus is concerned with sensibility of the opposite eyeball and its movements. On this lesser issue, Schäfer agreed with Ferrier. But our experiments on the occipital lobe have yielded no less definite positive results than those upon the angular gyrus yielded negative. I fail to understand how Dr Ferrier was unable ever to succeed in getting movements of the eyes by applying electrical stimulation to the occipital lobe ... removal [produced] immediate bilateral homonymous hemianopsia ... bilateral removal with the angular gyri quite intact [resulted] in total and permanent blindness ... the [reason why] Ferrier and Yeo failed to obtain any symptoms of defective visual perception [is that] the removal was very incomplete ... this also applies in some measure to the experiments of Mr Horsley and myself ... but when a small portion of one of the lobes is left ... the limit of the visual field of the retinae may be greatly restricted.
This analysis of the cortical representation of vision inflamed Ferrier, but it was in Schäfer's more extensive explorations of functional localization in the cerebral hemispheres that he offered something of a hostage to fortune: respecting the sense of hearing, the opinion that has obtained most currency is that of Dr Ferrier, to the effect that it is entirely localised in the superior temporal gyrus: but, when we come to sift the evidence ... we find it to be very insufficient ... in one monkey ... we destroyed the superior temporal gyrus upon both sides ... so that not a trace of the convolution could remain ... hearing was not perceptibly affected ... one monkey had the whole temporal lobe removed on both sides ... as far as the hippocampal margin ... this was marked by idiocy ... loss of intelligence and memory ... so that the animal received and responded to impressions from all the senses but appeared to understand very imperfectly the meaning of such impressions ... this was most evident for visual impressions ... but there was never any difficulty in observing that sounds were heard ... [and] Dr Ferrier's surmise that the antero-inferior extremity of the temporal lobe is related to taste ... is based on entirely insufficient evidence.
Ferrier reacted without restraint. Commenting at length on the January 1888 article, he rallied forth: Professor Schäfer ... who has done some good work in cerebral localisation ... not only differs in important respects from all who have preceded him ... but exhibits the curious spectacle of one who ... while he thus abounds in negations ... gives us no clue to the situation of the centres whose seat he so ruthlessly demolishes ... except that its removal ... is in some way the cause of temporary idiocy ... to one who has accepted the principle of localisation in respect to vision, tactile sensibility, and special motor faculties ... but [whose] assertion ... that hearing, taste and smell have no local habitation in the brain ... this appears a most lame and impotent conclusion. Having thus damned with faint praise, Ferrier felt free to correct any impression he might have given that he did in fact rate Schäfer's earlier work, reminding readers that, on a previous occasion: I gathered from Professor Schäfer that he had obtained evidence of an auditory centre in the [superior temporal gyrus] ... [he] is entitlednay boundto change his views when the evidence on which he based them appears [to him] on further consideration to be insufficient ... Professors Schäfer and Horsley gave me an opportunity of testing, and I am happy to say of correcting, conclusions which they were inclined to draw adverse to my own ... Professor Schäfer's figures (see Figures 14) must be regarded as a sort of pictorial representation of his own idea or wish ... rather than of reality. Clearly there had been an unpleasant exchange at a meeting of the Physiological Society on February 12, 1887, later repeated at the Neurological Society, and the rift was deepening. In the April issue, Ferrier drew attention to private correspondence such that the editor (Armand de Watteville) had to blow the whistle and inform readers that an objection raised by Professor Ferrier to the term "report" used in connection with an alleged Collective Report of a committee of the Neurological Society alluded to an individual expression of opinion by Drs Bastian, Brunton and Waller who took part in the investigation.
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Ferrier continued with a detailed discussion on who had said what with respect to the interpretation of pricking forward versus upwards of the auricle in the hearing monkey and the lop-eared rabbit, leading up to the reopening up of an old sorethe 1881 International Medical Congress, at which Ferrier had demonstrated Monkey F to the Physiological section and had a spat with Friedrich Goltz in claiming that it was possible to study cerebral localization by electrical stimulation without the need for ablation. In response to taunts from Goltz that dogs with virtually no remaining cerebrum could do everything attributed to Ferrier's sensory and motor centres, Ferrier pulled, as it were, a rabbit out of the hat by pointing out that a dog was not a monkey. The umpire was Sir William Gowers, who concluded that Goltz's dogs had rather more brain in situ than he had claimed. Incidentally, after the show Ferrier was prosecuted for cruelty to animals, but was acquitted. Clearly, Dr Ferrier was still smarting, since the April 1888 issue has five pages of notes describing Monkey F, the heroine of the 1881 International Medical Congress. (It was another of Ferrier's monkey's that elicited from Charcot the remark: it's a patient.)
After much airing of doubts and counter-arguments, Ferrier's position in April 1888 was that bilateral destruction of the upper two-thirds of the superior temporal gyrus produced deafness. I was not a little amused at listening to Professor Schäfer's comments on the above record ... that we had lighted on a specimen of a monkey stone-deaf before the operation ... it has been said that a dead donkey is difficult to find. A deaf monkey will be a greater curiosity. And when Professor Schäfer succeeds in finding one, it will not help his argument. There can be few more withering dismissals than Ferrier's next salvo: Professor Schäfer does not, in his paper, repeat (t)his brilliant inspiration, but, with a generosity which I do not fail to appreciate, sums up his estimate of the evidence that Professor Yeo and I [gave] ... with the declaration of his belief that he is justified in asserting that the supposed localisation of the auditory perceptive faculties in the temporal lobe in monkeys has no experimental evidence in its favour. Next, aiming to demolish Schäfer's failure to detect any disturbance of taste or olfaction after temporal lobe scooping, he continues: if Professor Schäfer is right in his conclusions that the whole of the temporal lobe, including the cornu Ammonis and the hippocampal lobule, may be removed without abolishing or impairing the sense of smell, he must ... consider that comparative anatomy is a delusion and a snare. But it was Schäfer's views on vision that so irked Ferrier, and to that he returned in closing; playing the advocate, he conceded the possibility, for the sake of argument, of several strands in Schäfer's case: the chief point at issue between him and myself is the anterior boundary of the visual zone ... on this the whole question turns. Pointing out that each of them had observed hemianopic defects following temporal lobe removal, and yet neither considered that to be the seat of the visual centrethe explanation being simply that the optic radiations had been damagedthe discrepancies in Schäfer's work were the result of his failure to distinguish tracts from the perceptual centre. Ferrier rested his case with: Professor Schäfer has furnished me with so many facts showing that the visual centres are not confined to the occipital lobes proper, but embrace also the angular gyri, that it seems unnecessary for me to insist further on the subject ... it is a pity that he did not examine his own facts a little more closely before committing himself ... while regretting for his own sake the attitude he has taken ... I cannot feel otherwise than indebted to him for inducing me to re-examine ... and re-affirm, with increased conviction, the views which I have previously enunciated.
On hearing, Ferrier has the edge; on taste and olfaction, the points are even; and on memory loss in bilateral temporal lobe lesions Schäfer was about 70 years ahead of the game. As for vision, both were constrained by the view that fibre tracts converge on a single structure specialized for perceptual appreciation; elucidation of the ocular dominance and orientation columns in VI (Brodmann area 17), the localization of V5 in the posterior bank of the superior temporal sulcus, and the recognition that vision is a complex percept with an equally complex functional anatomy, lay ahead (see page 1226). But if Schäfer and Ferrier worked on vision in the relative dark, this was clearly an era when the gloves were off with respect to freedom of speech, and editors felt no need to reign in the combatants.
Cambridge, UK
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