Special issue: Historical paperAlfred Walter Campbell and the visual functions of the occipital cortex
Introduction
Alfred Walter Campbell was an Australian neuropathologist whose pioneering cytoarchitectonic studies of the whole human brain were published between 1902 and 1905. In those works, Campbell differentiated two visual areas in the brain: one in which primary stimuli were registered and another where they were interpreted and given psychological meaning. Here I will suggest that Campbell's differentiation pointed to the areas we have come to term V1 and V2 and the way to differentiating other specialised visual areas. Although some aspects of Campbell's work on the visual system have been examined by ffychte and Catani (2005), I believe I provide a wider context and, in some respects, a more detailed scrutiny.
Campbell's identification of distinct areas of the brain by their cell and fibre structure was part of his study of areas that could be related to areas of known function. They were at once recognised internationally as providing the authoritative view of the brain's structure in relation to its functions. Yet, by the 1930s Campbell himself was virtually forgotten and his work had been overshadowed by Brodmann's similar but later mapping. As a consequence, his contribution to our understanding of vision has also been overlooked.
Here I provide some background to Campbell's work. After describing his early life and education in Australia, and his Scottish medical education and early work as a pathologist at the Lancashire County Lunatic Asylum at Rainhill near Liverpool, I summarise his work on the human brain. In describing the structures he identified in the occipital lobes, I analyse the similarities and differences between them and the related structures identified by others, especially by his critic Joseph Shaw Bolton.
I conclude by proposing some reasons for how Campbell's work came to be overshadowed by the later studies of Brodmann and for the more general lack of recognition given him and his work. Those reasons include the effect of the controversies precipitated by Campbell's alliance with Charles Sherrington over the functions of the sensory and motor cortices.
Section snippets
Family background, early life, and education
As his name suggests, Alfred Walter Campbell (Fig. 1) was of Scottish extraction but, unlike many Scottish migrants who settled in Australia, the Scots were not the only contributors to his lineage: Danes and French Huguenots were also significant.
Alfred Walter's great-grandfather, John Campbell, was a Scot who married Elisabeth Sophie Berg, in Copenhagen in 1796. By 1826 John was in Tharangambadi [Trankebar or Tranquebar], one of the Danish East India Company's colonies some 120 km south of
Medical education and ‘Mental Science’
We do not know why Alfred Walter chose medicine as a career but he must have decided on it during his ‘Oaklands’ years. Nor do we know why he chose Edinburgh. All we know is that he left Sydney in March 1885 to arrive in Britain probably some 7 weeks later and successfully pass the Edinburgh ‘Preliminary Examination for Degrees in Medicine’ in the August. The standard for this entrance examination was that of the degree of Bachelor of Arts. Edinburgh medical students then qualified to practice
Campbell's histopathological work at Rainhill
Apart from the time consuming work of conducting post-mortem examinations on each of the patients who died in Rainhill—about 10–12 per month—Campbell found time to continue the kind of research he had begun in Prague, to set up more adequate pathological facilities, to establish a museum of specimens relevant to mental illness, and to begin somewhat fateful collaborations with Henry Head on human dermatomes and Charles Sherrington on the localisation of motor function.
In fact it was literally
Rainhill, Campbell, and Henry Head's dermatomes
It is well known that Alfred Walter Campbell collaborated with Henry Head in histological research on herpes zoster (shingles) relating to his concept of dermatomes. It is commonly assumed that Head went to Rainhill for that purpose. Although it is true that he did take up a position of unpaid Clinical Clerk at Rainhill in December 1894 and that Campbell and he did eventually work together, the herpes work did not begin until the following year and the bulk of it was carried out by Campbell.
Campbell's identification of the visual areas in the human and animal brain
Towards the end of the 1890s Alfred Walter Campbell began to extend his histopathological studies into topics not directly related to mental illness. The new direction is evident in his using Marchi staining to investigate the tracts of the spinal cord and in his developing a method using osmic acid to stain medullated fibres (Campbell, 1897a,b).
We do not know why Campbell made this extension but it may have been partly due to the appointment in 1895 of Charles Scott Sherrington [1857–1952] to
Campbell and the motor systems
The visual system was not Campbell's primary concern. Rather it was almost an unrelated accompaniment to his seeking a histological basis for the motor area identified in the physiological experiments of Sherrington and Grünbaum at the University of Liverpool. Sherrington and Grünbaum used faradic stimulation to localise motor function in the cerebral cortices of several chimpanzees, an orangutan, and a gorilla. They found that only stimulation in front of the central sulcus, or fissure of
Campbell's identification of structures in the human and animal brain
Campbell's localisation work had begun with the investigation of nerve fibres, and in a single chimpanzee. It soon became a more ambitious project aiming to test whether all the known functional areas of the human and animal brain could be identified by differences in the layering (lamination) of their cells as well as the nature of their fibres. He reported the results of this larger endeavour on 20 June 1903, published part of them in July 1904, and in complete detail in 1905 (Campbell, 1903a
Campbell's identification of the visuosensory and visuopsychic areas
Sherrington communicated Campbell's first report containing his new cell and fibre based mappings of the human brain to the Royal Society on 17 November 1903 where it was read on 3 December. On 8 August the following year Campbell communicated, again through Sherrington, a second part on the cat, dog, and pig. It was read on 1 December 1904 (Campbell, 1903–1904, 1904–1905). Both Royal Society referees, Frederick Walker Mott [1853–1926] the English neuropathologist and psychiatrist, and William
Campbell's identification of the motor areas
Just as Bolton's visuosensory cortex was located in the same place as Campbell's, so the motor cortex defined physiologically by Sherrington and Grünbaum was matched by Campbell's precentral and intermediate precentral areas (Fig. 8). Both were anterior to the fissure of Rolando and different histologically from the two areas immediately behind it. How Campbell differentiated them is illustrated in Fig. 11. The precentral fibre representation was, as he had said in 1902, generally superior to
Controversies about Campbell's localisation of the visual and motor areas
Despite the favourable reception given Campbell's work, there was some hostility, partly over his delineation of the visual and motor areas and partly because he had attributed functions to them. Bolton launched an attack on his visual work because he believed Campbell was claiming priority in identifying the visuosensory area but was mostly concerned at what he took to be Campbell's denigration of his work on the frontal lobes. A more vicious attack, one with serious consequences for Campbell,
Campbell's work: forgotten and overshadowed
More than the passing of time has ensured that many of Alfred Walter Campbell's pioneering discoveries are now forgotten. To that passage must be added the effects of the criticism, misrepresentation, and the deliberate disregard of his work by important figures in English neurology and neurosurgery. Campbell's cytoarchitectonic work including his recognition of the two areas of the occipital cortex suffered particularly.
Campbell's well-known modesty also contributed—apart from his brief reply
The return to Australia
Alfred Walter Campbell returned to Australia in December 1905 and it is something of a mystery why he did so and why it was at that particular time in his career. Prof. Mervyn Eadie, the historian of Australian neurology, who was fortunate enough to be able to interview Campbell's daughter, Mrs. Veda Hope, as well as a number of Campbell's colleagues, believes that Campbell's decision to marry was among the factors that led to his returning (Eadie, 2000, pp. 45, 53, 2001). Walterina Jean
Campbell's achievements
Campbell married within months of returning and became Australia's first neurologist. He did some quite important but little noticed research on localisation in the cerebellum (Campbell, 1913), the brain of the gorilla (Campbell, 1916a), and, based largely on his Australian Army Medical Corps experience of the effects of the Gallipoli campaign, published on neuroses and psychoses in war (Campbell, 1916b). As World War I was ending, he joined with John Burton Cleland [1878–1971], Principal
Acknowledgements
I should like to thank Stephanie Forkel of London and Ian Macmillan of Palmerston North, New Zealand, for their considerable help with the figures and the two anonymous reviewers whose sharp comments considerably improved the first version of this paper.
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