CEPA eprint 2927

Integrating Mind and Brain: Warren S. McCulloch, Cerebral Localization, and Experimental Epistemology

Abraham T. H. (2003) Integrating Mind and Brain: Warren S. McCulloch, Cerebral Localization, and Experimental Epistemology. Endeavour 27(1): 32–38. Available at http://cepa.info/2927
Table of Contents
McCulloch’s early years
J.G. Dusser de Barenne, Rudolf Magnus and the physiological a priori
Strychnine and functional localization
Recently, historians have focused on Warren S. McCul­loch’s role in the cybernetics movement during the 1940s and 1950s, and his contributions to the develop­ment of computer science and communication theory. What has received less attention is McCulloch’s early work in neurophysiology, and its relationship to his philosophical quest for an ’experimental epistemology’ – a physiological theory of knowledge. McCulloch’s early laboratory work during the 1930s addressed the problem of cerebral localization: localizing aspects of behaviour in the cerebral cortex of the brain. Most of this research was done with the Dutch neurophysiolo­gist J.G. Dusser de Barenne at Yale University. The con­nection between McCulloch’s philosophical interests and his experimental work can be expressed as a search for a physiological a priori, an integrated mechanism of sensation.
Through his quest for an ’experimental epistemology’ – a physiological theory of knowledge – neurophysiologist Warren Sturgis McCulloch (1898-1969) aimed to discover how the brain functions to sustain mental phenomena such as sensation and perception. Integrating knowledge of behaviour – functions of the mind – to neurological processes in the brain, McCulloch asked questions that were at once physiological, psychological and philosophi­cal. Recently, historians have emphasized McCulloch as a key player at the Macy Conferences on Cybernetics during the late 1940s and 1950s, and as an important figure in the development of computer science and theories of infor­mation processing [1-3]. What has received less attention is McCulloch’s early work as a neurophysiologist strongly motivated by philosophical and epistemological concerns.
Much of McCulloch’s experimental research on living systems was done during his medical and psychiatric training in the New York City area during the 1920s and during his fruitful collaboration with the neurophysiolo­gist J.G. Dusser de Barenne (1885-1940) at Yale Univer­sity during the 1930s. Within these clinical and laboratory settings, McCulloch’s work involved ’localizing’ aspects of behaviour in the cerebral cortex – the postulated ’material source’ of psychological functions in the brain. As Robert Young observed in his classic account of brain research in the 19th century, cerebral localization can be seen to fall within the ’empirical and conceptual domain within which the mind-body problem was – and continues to be – investigated’ [4]. When viewed in this light, McCulloch’s work on cerebral localization enabled him to pursue his philosophical quest – integrating mind and brain – in a scientific and experimental setting. This essay will focus on McCulloch’s work on cerebral localization, particularly his research with Dusser de Barenne, with the aim of shedding light on the connections between his experimen­tal work and his philosophical interests.
McCulloch’s early years
Warren McCulloch received his bachelor’s degree from Yale University in 1921, with a major in philosophy and a minor in psychology. Here he became immersed in the philosophy oflmmanuel Kant, and began formulating questions on the relationship between knowledge, thought and the physiology of the brain. McCulloch received his M.A. in psychology from Columbia University in 1923, and eventually went on to medical school at Columbia, receiving his M.D. in 1927. In 1928, McCulloch interned as a neurologist under Robert Foster Kennedy (1884-1952) at Bellevue Hospital in New York City, doing experimental research on epilepsy and head injuries.
From 1929 to 1931, McCulloch taught physiological psychology at Columbia’s newly created Seth Low Junior College in Brooklyn, and published several papers with neurologist Frank H. Pike (1876-1953), who had been his mentor in the Department of Physiology at the College of Physicians and Surgeons at Columbia University [5]. Their focus was on the localization of areas of the cortex associated with epileptic convulsions [6], [7]. Localization, in this clinical setting, was seen to bear on the problem of diagnosis – for instance, if a cyst was suspected to be associated with symptoms of convulsive movement, knowl­edge of its location in the brain and of the cortical areas thought to be involved in convulsions would facilitate diagnosis. In a broader sense, Pike and his colleagues also saw the problem of localizing cortical areas involved in convulsions as a special case of the functional organization of the motor system, and of the functional localization of nervous mechanisms in general. McCulloch’s experiments with Pike primarily involved inducing convulsions in cats, usually through an intravenous injection of a solution of absinthe. Pike, McCulloch and their colleagues attempted to determine the area of the cortex involved in the convulsive movements by making a lesion of part of the motor cortex and then administering the convulsion-producing agent. In the first few days after the infliction of lesions, only mild seizures were observed after injection of absinthe.
Motivated by the observation of British neurosurgeon, John Hughlings Jackson (1835-1911), that a remarkable degree of recovery could be observed in patients several weeks or months after removal of the motor cortex [8], Pike and his colleagues performed absinthe experiments on animals whose cortical areas had been excised between several months and two years before the experiment. Following this, more severe seizures could be induced by absinthe, indicating a recovery of motor functions despite removal of areas of the motor cortex. They interpreted their results as providing evidence for Jackson’s idea of an ’integration’ between ’higher’ and ’lower’ motor areas.
Between 1932 and 1934, in the depths of the Depression, McCulloch took a series of appointments – primarily to earn money – at the Rockland State Hospital for the Insane in Orangeburg, New York. In 1934 he returned to Yale University as an Honorary Research Fellow in Dusser de Barenne’s Laboratory of Neurophysiology, who was to be his mentor and collaborator for the next six years.
J.G. Dusser de Barenne, Rudolf Magnus and the physiological a priori
Johannes Gregorius Dusser de Barenne began investigating the physiological effects of strychnine on spinal cord reflexes whilst a teaching assistant at the Laboratory of Physiology at the University of Amsterdam from 1909 to 1911 [9]. Whilst serving as a medical officer in the Dutch Army at Delft during the First World War, Dusser de Barenne’s attention turned to the functional localization of sensory phenomena in the cerebral cortex.
Dusser de Barenne’s early work using the strychnine method on the cortex was done during his period at Delft, and involved experiments on the cerebral cortex of the cat [10]. After anesthetizing the animal with chloroform and ether, the region of the cortex to be experimented on was exposed, and any excess cerebrospinal fluid was absorbed by dabbing the surface of the cortex with cotton. A 1 % strychnine solution, coloured with toluidin blue, was then applied to the cortex using a tiny wad of cotton wool at the end of forceps. The resulting poisoned spot on the cortex was then seen as a small blue area of a few square millimetres. The cat’s skin was then stitched back, to prevent cooling. Following the animal’s recovery from narcosis, Dusser de Barenne would observe and compare symptoms when the sensory cortex was strychninized within a certain region of the cerebral cortex and then outside this same region, observing behavioural disturb­ances in the cat, such as paralysis and hypersensitivity.
Just before the end of the war, Dusser de Barenne left the army to take up an appointment in the Departments of Pharmacology and Physiology at the Utrecht University. Here he became an assistant to the eminent German neurophysiologist Rudolf Magnus (1873-1927), and together they studied the physiology of posture [11]. Magnus had been chair of pharmacology at Utrecht since 1908, and although he was a successful pharmacol­ogist, he became better known for his work in neuro­physiology, particularly on the physiology of posture and reflex mechanisms. In addition to his neurophysiological studies, Magnus was also known for his notion of the ’physiological a priori’. In a lecture delivered at Stanford University in 1927, Magnus pointed to Kant’s demon­stration that in everything we know – our observations and conclusions about the world around us – there are ’numerous elements, which are given a priori, and which we are therefore compelled to employ in any experience in thinking and drawing our conclusions’ [12]. Magnus argued that Kant’s synthetic a priori should be interpreted not only ’philosophically-psychologically’, as had been done in the past, but also physiologically – that part of these a priori factors in our psyche must have a physiological basis. This was related to the notion that one does not come to sensory data as a ’blank tablet’, but rather brings a sort of relational structure within the nervous system to interpret sense data. The nature of our sensory impressions is determined a priori, Magnus argued, by the physiological apparatus of our senses: the sensory nerve centers in the brain. The localization of these sensory areas in the brain became the focus of Dusser de Barenne’s research.
In the spring of 1924, Dusser de Barenne went to the laboratory of the eminent neurophysiologist Charles Scott Sherrington (1857-1952) at Oxford University. There, he studied sensory symptoms through the application of strychnine to the cerebral cortex of rhesus (macaque) monkeys, and produced results that delimited the sensory cortex of the monkey [13]. In 1929, Dusser de Barenne was recruited by Yale University’s Dean of Medicine, Milton C. Winternitz, to join the Yale faculty. In the fall of 1930, Dusser de Barenne arrived at Yale and was made Sterling Professor of Physiology, and eventually built the Labora­tory of Neurophysiology at the School of Medicine. Here he continued his work on the localization of sensory functions in the cortex using the strychnine method.
Strychnine and functional localization
During the first decades of the 20th century, two methods of cerebral localization were prominent: the ’lesion’ or ’extirpation’ method (as used in Pike’s laboratory at Columbia), and the method of electrical stimulation. The lesion method involved cutting and removing areas of the cortex (grey matter) and then observation of the animal’s behaviour and motor functions. The loss of tissue in specific areas of the cortex was then related to the alteration of the animal’s behaviour. In the method of electrical stimulation, different parts of the cortex and peripheral nerves were subjected to an electrical current, and sensory and motor functions were mapped onto the brain depending on the location of stimulation [14].
In his Ferrier Lecture of 1938, the Nobel Prize-winning neurophysiologist Edgar D. Adrian (1889-1977) observed that up until then, the electrical stimulation of the cortex had been seen as useful for revealing the ’receiving and executive apparatus of the cerebrum but can show very little of the processes which intervene between the incoming and outgoing signals’ [15]. In a sense, he argued, we only have a mechanism of ’entrances and exits’. This was the tradition that had dominated studies of the cerebral cortex since Eduard Hitzig’s (1838-1907) and Gustav Fritsch’s (1837-1927) pioneering work on the electrical activity of the brain and their demonstration of the localization offunction in the monkey [16]. For Adrian, the problem with simply making electrical recordings from the cortex after sensory stimulation was that there was no indication of what the records represented in terms of cell activities and pathways. In his review of recent work, Adrian noted that Dusser de Barenne’s group at Yale had been making headway in analysing these electrical records – through the application of strychnine – and thus were contributing to the understanding of the organization of activity within the cortex.
Most of the work Adrian referred to here were the results of Dusser de Barenne’s work with McCulloch. Between 1934 and 1940, they published over 25 papers together, the bulk of which outlined experiments using strychnine as a tool for localization. Methodologically speaking, Dusser de Barenne felt that strychnine was a more precise tool for determining the complexities of the functional organization of the cortex. For Dusser de Barenne, the standard method of stimulation had yielded no significant results relevant to the problem oflocalizing sensory functions in the cortex. Dusser de Barenne also saw several advantages of his strychnine method over the extirpation method. Firstly, the application of strychnine to the cortex caused symptoms of excitation with more precision and certainty, and allowed an easier interpret­ation of symptoms. By contrast, the extirpation method only resulted in symptoms of impairment of sensation, which were often vague. The strychnine method, he argued, was also simpler – it was less invasive, and induced less stress in the animal under study, whilst the extirpation method was often accompanied by additional pathological changes owing to shock and trauma, which confused results. Unlike the extirpation method, the strychnine method could be used without damage to the brain, and could produce precise results with relatively short experiments [17].
Dusser de Barenne was critical of’classical’ localization theory, with its assumption of a ’sharp, point to point, geometrical projection of the body on the cortex’ [18]. For example, it is incorrect to say that a sensory function such as vision is ’localized’ in the cerebral cortex, for this function also involves concomitant activi­ties at other levels of the nervous system. Dusser de Barenne argued that the term ’localization of function in the cerebral cortex’ should be replaced by the term ’functional localization in the cerebral cortex’.
This conceptual framework informed Dusser de Barenne’s research with McCulloch. McCulloch’s earlier work with Pike in a clinical setting had convinced him that the idea of strict functional localization in the cortex was flawed. Rather than simply localizing sensory functions, Dusser de Barenne and McCulloch aimed to determine how these functions might work together in the process of sensation, in a sense, elucidating a physiological a priori.
Early on after McCulloch’s arrival, Dusser de Barenne’s lab had been using various methods to investigate the electrical activity of the cortex, and sensory stimulation in particular. Eventually Dusser de Barenne’s strychnine method was used to investigate the functional organiz­ation of the sensory cortex, and its relation to other areas, particularly the thalamus. In 1936, Dusser de Barenne and McCulloch published their first results employing the strychnine method [20]. Here, through the coupling of the strychnine method with the recording of electrical activity of the cortex, they established that there were functional boundaries between the main subdivisions of the sensory­motor cortex. They observed that the application of strychnine was accompanied by large, rapid changes in the action potentials (which they termed ’strychnine spikes’) recorded from specific areas of the cortex; and that the spikes were dissimilar when recorded from different areas of the cortex. Dusser de Barenne and McCulloch also found that strychninization of a small area in one of the subdivisions of the sensory cortex, for example the arm area, could result in changes in action potentials of the whole subdivision. Furthermore, the character of these spikes was specific for each area strychninized. And finally, their work revealed that there were directed functional relations between areas: for example, they observed that if region A was strychninized, spikes were recorded from region B, but if region B was strychninized in a separate experiment, no spikes were recorded from region A. Their work confirmed Dusser de Barenne’s earlier hypothesis that complex functional relationships exist between different areas of the cortex [21-23]. Dusser de Barenne and McCulloch concluded that for certain areas of the cortex, the effects of strychnine superseded structural boundaries of the cortex but respected func­tional boundaries.
Dusser de Barenne and McCulloch also used the strychnine method for delimiting neurons in the cerebral cortex, a procedure called ’chemical neuronography’, to study the interaction of various cortical and subcortical regions of the brain [24]. Their method here was similar to that used in their earlier work; however, their goal here was to understand communication in the cortex by deducing specific pathways of neural impulses through an analysis of their experimental results. Based on their previous work mapping functional areas in the cerebral cortex, Dusser de Barenne and McCulloch aimed to correlate these findings with the neuronal structure of the cortex. Their goal was to determine if there were individual neurons in the area of the cortex where strychnine was applied that had ’endings’ in the area of the cortex where the electrical activity was being recorded. Drawing on some anatomical evidence regarding the direction of neuronal connections in the cortex, Dusser de Barenne and McCulloch observed that when one strychninized a particular region A, and recorded ’spikes’ from region B, the neurons that were strychninized in region A had an ending in region B. They concluded that local strychninization of the cerebral cortex coupled with the electrical recording was a powerful tool for delimiting the origin and ending of neurons in the central nervous system.
McCulloch continued to collaborate with Dusser de Barenne for six years. Their last joint publication, on the sensory cortex of the chimpanzee, appeared in 1940 [25]. Here they collaborated with Percival Bailey (1892-1973), who was then on leave from the Department of Neurology and Neurosurgery at the University of Illinois. In 1941, after Dusser de Barenne’s death the previous year, McCulloch, along with others from Dusser de Barenne’s lab at Yale, was invited by Bailey to join the Illinois Neuropsychiatric Institute (NPI) in Chicago. Over the next several years, McCulloch and his colleagues continued using the strychnine method to study the functional organization of the sensory cortex and motor cortex of the monkey and chimpanzee.
McCulloch’s early training in philosophy, psychology and physiology brought about a mingling of intellectual pursuits. From an early stage in his career, McCulloch was directed by a search for an experimental epistemology – in his words, a physiological theory of knowledge [26], [27]. He was drawn into neurophysiology, in a sense, by a philosophical problem. His search for a physiological basis of knowledge acquisition was in essence a quest to understand the neurophysiological basis of sensation and perception, a goal he pursued during his years of collaboration with Dusser de Barenne. Key to McCulloch’s work with Dusser de Barenne was an interest in the organization and integration of the nervous system, specifically, in the cerebral cortex. In their strychnine experiments, they were investigating not only the local­ization of sensory functions in the cortex, but how the system of neuronal connections within the cortex worked as an integrated whole. For McCulloch, it was this organization and connectivity of the sensory cortex in the brain embodied Kant’s a priori element – the physiological a priori was, for McCulloch, an integrated mechanism of sensation. Thus, in his work with Dusser de Barenne, McCulloch was able to pursue his quest for an experimental epistemology, and relate the psychological functions of sensation and perception to the neurophysiol­ogy of the brain.
[1] Kay, L.E. (2001) From logical neurons to poetic embodiments of mind: Warren S. McCulloch’s project in neuroscience, Science in Context 14 591-614
[2] Heims, S.J. (1993) Constructing a Social Science for Postwar America: The Cybernetics Group, MIT Press, pp. 1946-1953
[3] Edwards, P.N. (1996) The Closed World: Computers and the Politics of Discourse in Cold War America, MIT Press
[4] Young, R.M. (1990 1970) Mind, Brain, and Adaptation in the Nineteenth Century, p. vii, Clarendon Press
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[6] Pike, F.H. et al. (1929) Some observations on experimentally-induced convulsions, Am. J. Psych. 9 259-283
[7] Pike, F.H. et al. (1930) The problem oflocalization in experimentally induced convulsions, Archives of Neurology and Psychiatry (Chicago) 23 847-868
[8] Jackson was known for his clinical research on epileptic patients and for his idea of a ’hierarchy’ of brain functions. Arguing against the idea of a strict localization of function in the cortex, Jackson purported that a change in the quantity of ’nervous energy’ that flowed through a pathway in the cortex occurs following injury to another pathway, and that the intact pathway as a result had a change in its functional capacity. For an examination of Jackson’s work in the context of late 19th-century brain research, see Harrington, A. (1987) Medicine, Mind, and the Double Brain, Chapter 7, Princeton University Press
[9] Fulton, J.F. and Gerard, R.W. (1940) J.G. Dusser de Barenne 1885- 1940, J. Neurophysiol. 3 283-292
[10] Dusser de Barenne, J.G. (1916) Experimental researches on sensory localisations in the cerebral cortex, Quarterly Journal of Experimental Physiology 9 355-390
[11] Magnus, R. and Dusser de Barenne, J.G. (1920) Beitrii.ge zum Probleme der Korperstellung. III. Die Stellreflexe bei der grosshirn­losen Katze und dem grosshirnlosen Runde, Pflugers Archiv fiir gesamte Physiologie 180 75-89
[12] Magnus, R. (1930) The physiological a priori, Lane lectures on experimental pharmacology and medicine, Stanford University Publi­cations, University Series, Medical Sciences. (vol. 2, no. 3) pp. 97-103, Stanford: Stanford University Press
[13] Dusser de Barenne, J.G. (1924) Experimental researches on sensory localization in the cerebral cortex of the monkey (Macacus), Proc. R. Soc. London, Series B 96 272-291
[14] For the development of electrical instrumentation in studies of the brain, see Brazier, M.A.B. (1961)A History of the Electrical Activity of the Brain, Pitman
[15] Adrian, E.D. (1939) Ferrier Lecture: the localization of activity in the brain, Proc. R. Soc. London, Series B 126 433-449
[16] Fritsch, G. and Hitzig, E. (1870) Uber die elektrische Erregbarkeit des Grosshirns. Archiv fiir Anatomie, Physiologie und wissenschaftliche Medicin 300-332. Reprinted in Von Bonin, G. (trans, ed.) (1960) Some Papers on the Cerebral Cortex, pp. 73-96, Charles C. Thomas
[17] Dusser de Barenne, J.G. (1916) Experimental researches on sensory localisations in the cerebral cortex, Quart. J. Exp. Physiol. 9 355-390 18 Dusser de Barenne, J.G. (1933) ’Corticalization’ of function and functional localization in the cerebral cortex, Archives of Neurology and Psychiatry (Chicago) 30 884-901. p. 894
[19] Dusser de Barenne, J.G. (1935) A word of criticism on the designation ’localization’ of function in the cerebral cortex, Archives of Neurology and Psychiatry (Chicago) 33 1081
[20] Dusser de Barenne, J.G. and McCulloch, W.S. (1936) Functional boundaries in the sensori-motor cortex of the monkey, Proc. Soc. Exp. Biol. Med. 35 329-331
[21] Dusser de Barenne, J.G. and McCulloch, W.S. (1936) Some effects of local strychninization on action potentials of the cerebral cortex of the monkey, Trans. Am. Neural. Assoc. 1936 171 62nd Annual Meeting, Atlantic City, NJ, June 1-3, 1936.
[22] Dusser de Barenne, J.G. and McCulloch, W.S. (1938) Functional organization in the sensory cortex of the monkey (Macaca mulatta), J. Neurophysiol. 1 69-85
[23] Dusser de Barenne, J.G. and McCulloch, W.S. (1938) The direct functional interrelation of sensory cortex and optic thalamus, J. Neurophysiol. 1176-186
[24] Dusser de Barenne, J.G. and McCulloch, W.S. (1939) Physiological delimitation of neurones in the central nervous system,Am. J. Physiol. 127 620-628
[25] Bailey, P. et al. (1940) Sensory cortex of the chimpanzee, J. Neurophysiol. 3 469-485
[26] McCulloch, W.S. (1989) Recollections of the many sources of cybernetics in The Collected Works of Warren S. McCulloch (McCul­loch, R., ed.), (vol. I, pp. 21-49), Intersystems. Reproduced fromASC Forum (1974), vol. 6, pp. 5-16)
[27] McCulloch, W.S. (1964) A historical introduction to the postulational foundations of experimental epistemology, in: F.S.C. Northrop (Ed.), Cross-Cultural Understanding: Epistemology in Anthropology, Harper &Row,pp. 180-193
[28] Marshall, L.H. and Magoun, H.W. (1998) Discoveries in the Human Brain: Neuroscience Prehistory, Brain Structure, and Function, Humana, p. 84
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