SEPTEMBER

1963

BRAIN
VOL. 86, PART 3. SOME REFLECTIONS ON BRAIN AND MIND 1
BY

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THE ever-growing number of articles, books and symposia devoted to one or other aspect of the brain-mind relationship is a remarkable feature of our time. Since it has been debated since the time of the Greeks, and the main questions about it were clear in general at least three hundred years ago, it is perhaps natural to ask why so much attention should be devoted to it today. When men knew only the most general facts about the relationship of the brain and the mind, the problem of its nature necessarily presented itself in the most general terms, as a philosophical problem involving supposed mental entities with certain characteristics, and matter in general, since of the material structure of the brain virtually nothing was known. It is the vast expansion of our knowledge of neurophysiology and psychophysiology which has stimulated the current surge of interest in the brain-mind relationship. Two main questions are asked about it, and it is important to keep them distinct, at least initially. Most people are aware that correlations exist between the brain and the mind,8 and the observations of these is part of the everyday work of the neurologist. If we want to know what these correlations are, we must investigate, describe and clarify them by every available method. But we may also ask what the nature of the correlation of brain and mind is in general. It is widely believed that these two lines of enquiry have no relation to each other. Hughlings Jackson (1931, 1, 367)* was concerned solely with the first question. "States of consciousness," he wrote, "are assumed . . . to be merely concomitant with certain nervous states, those of the highest cerebral centres. I have nothing whatever to say of the
Based upon the Hughlings Jackson lecture delivered to the Montreal Neurological Institute, May 9, 1963. I purposely do not define these terms: "brain" is self-explanatory, and I here use "mind" in the everyday sense in which ideas and feelings for example are said to be "in the mind," or mental. Throughout thispaper the Hughlings Jackson references are to the Selected Writings.
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nature of the relation of the two utterly different and yet concomitant things, cerebration and mentation, to one another. As an evolutionist I am not concerned with this question and for medical purposes I do not care about it." Some contemporary workers think that those who study the brain should be equally cautious about discussing it. Granit (1955), in his book, Receptors and Sensory Perception, says: "I shall restrict my comments on epistemology to stating that the time is past when physiologists ex cathedra could issue naive views on the nature of knowledge, 'brain and mind,' reality and appearance, and similar concepts without penetrating the philosophical aspects of these problems in detail. They will merely be snared by their own lack of insight in the logical analysis of the conventions of language." But most of that seems to be equally true if repeated with the interchange of two words. The time is past when philosophers ex cathedra could issue naive views on the nature of knowledge, "brain and mind," reality and appearance, and similar concepts without penetrating the physiological aspects of these problems in detail. I do not expect that this question, one of the most fundamental problems of life, will be rapidly clarified, but I do not think it will take the four hundred years which Sherrington once predicted in conversation with me. The better we understand the correlation between the brain and the mind, and how the brain subserves mental functions, the more likely, I believe, shall we be to understand the nature of the relations between the two. So I am going to begin by discussing some aspects of the physiology of perception. This will be followed by a consideration of time in relation to the nervous system and the mind. That will lead to a discussion of the nature of consciousness and its physiological basis, and, finally, I shall say something about the relationship between brain and mind. My object throughout is not to present conclusions, but to ask questions to which I hope time will provide the answers.
RECENT ADVANCES IN THE PHYSIOLOGY OF PERCEPTION

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The newer electrophysiological methods are adding fundamentally to our knowledge of the physiological basis of perception. Micro-electrodes make it possible to record from single units at different levels of the nervous system, and so to discover how these respond to different kinds of sensory stimulus. For some years now it has been known that in the retina and optic nerve there are some units which respond to the onset of illumination, and some which respond to its cessation, and others yet again which respond to both. This is known as the on/off system (fig. 1). Hubel (1959) by means of implanted micro-electrodes has studied the activity of single units in the striate cortex of unrestrained cats. He observed that most cortical units showed no response to a mere change of background illumination. Many responded to a stationary light source

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FIG. 1.Diagram illustrating three fibres in the optic nerve firing spontaneously and their responses to illumination. 1, The on response; 2a, the off response; 2b, the off response inhibited by illumination; 3, a unit showing both on and off responses. (From "Receptors and Sensory Perception" by Ragnar Granit (1955).)

by showing on-and-off type responses (fig. 2). A response to movement was also observed. Some units responded to movement in one horizontal direction, but not to that in the opposite. In the same cerebral hemisphere Hubel observed two types of unit, one responding to leftward and the other to rightward movement. He also noted that movement could activate a single cortical unit, when it occurred over a relatively large region of the visual fieldof the order of 20, equivalent to 5 mm. on the retina.1 This observation indicated a convergence from a considerable retinal field upon a single cortical unit. Hubel concluded that "the striate cortex is not simply a retinotopically organized point-to-point relay station along the visual pathway." Rather it "would seem to contain units with receptive fields of widely differing extent." Hubel and Wiesel (1959) working with acute cat preparations also observed that most units in the visual cortex corresponded to a restricted retinal area, and that most such retinal fields were divided into excitatory and inhibitory regions, which were mutually antagonistic in relation to cortical units (figs. 3 to 5). They concluded that "the phenomena of summation and antagonism within receptive fields seem to provide a basis for the specificity of stimuli, in shape, size and orientation." Studies of binocularly activated units showed that the receptive fields in the two eyes were alike. The effect of this would be that the retinal images of objects behind, or in
Thus physiology has confirmed observations made by Riddoch (1917) during the first world war that after an occipital injury awareness of movement might recover before recognition of an object, thus demonstrating in man "that movement should be given a place among stimuli which are recognized as originating visual perceptions."
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Fio. 2.Responses of a cell in the cat's striate cortex to a 1 spot of light. Receptive field located in the eye contralateral to the hemisphere from which the unit was recorded, close to and below the area centralis, just nasal to the horizontal meridian. No response evoked from the ipsilateral eye. The complete map of the receptive field is shown to the right, x, areas giving excitation; A . areas giving inhibitory effects. Scale, 4. Axes of this diagram are reproduced on left of each record, a, 1 (0 -25 mm.) spot shone in the centre of the field; b-e, 1 spot shone on four points equidistant from centre; f, 5 spot covering the entire field. Background illumination 0-17 log. m.c. Stimulus intensity 1-65 log. m.c. Duration of each stimulus 1 sec. Positive deflexions upwards (Hubel and Wiesel, 1959).

front of, the point of visual fixation would not fall on corresponding parts of the visual fields, and their effects would not necessarily sum. They might antagonize each other, or not interact at all. This, the authors think, may produce an increased awareness of objects, and cause those at the same distance as the object fixed to stand out in relief. Hubel and Wiesel (1960) also studied the receptive fields corresponding to optic nerve fibres in the spider monkey, and obtained similar results to those in the cat. They demonstrated the existence of concentric retinal fields with on-and-off centres, either of which might lie outside the other. The same authors (Hubel and Wiesel, 1961) were able to demonstrate the same responses in the lateral geniculate body of the cat, the organization of which, they found, resembled that of the retina more than that of the striate cortex. In later papers these authors discuss their observations in relation to the architecture of the cat's visual cortex (Hubel and Wiesel, 1962, 1963; Hubel, 1963).

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Fio. 3.Responses of a unit to stimulation with circular spots of light. Receptive field located in area centralis of contralateral eye. (This unit could also be activated by the ipsUateral eye.) a, 1 spot in the centre region; b, same spot displaced 3 to the right; c, 8 spot covering entire receptive field. Stimulus and background intensities and conventions as in fig. 2. Scale, 6. x, areas giving excitation; A areas giving inhibitory effects (Hubel and Wiesel, 1959).

The work of Maturana, Lettvin, McCulloch and Pitts (1960) on the anatomy and physiology of vision in the frog has shown that the ganglion cells of the frog's retina form five natural classes. Class 1 respond to a sharp edge; class 2 to a convex edge; class 3 to changing contrast; class 4 to dimming; and class 5 measure the light intensity. The first four classes "act on the visual image to perform complex analytical operations that remain invariant under changes of the general outlook of the visible environment." "The activity of each ganglion cell is the result of the activity in a whole neighborhood and measures a particular quality in that neighborhood of numerous receptors" and, "because of the great overlapping of the dendritic arbors, numerous different ganglion cells are performing their corresponding operations on the same visual neighborhood." The projection of the retina on the tectum "provides that each class of ganglion cells has unique relationships with the superficial tectal neuropil." Hence "there is in the tectum a functional and spatial representation of the visible world of the frog." The movement of a fly, the frog's prey, is therefore signalled to it by the information derived from the response of specific ganglion cells to a highly specific stimulus. The image of an object presents itself to the retina as a matrix of discrete points. The activity of the retinal receptors and ganglion cells transforms this "into a matrix of overlapping contexts," which "establishes the systems of finite differencing in space whereby local continuity and curvature of boundaries are taken, thus giving rise at the same time to the notion of local extension," and, of course, movement, so that instead of a discrete point-space what is presented to the brain is information

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FIG. 4.Responses evoked only from contralateral eye. Receptivefieldjust outside nasal border of area centralis. a, 1 spot covering the inhibitory region; b, right half of a circle 12 in diameter; c, light spot covering regions illuminated in a and b. Background and stimulus intensities and conventions as in fig. 2. Scale, 12 (Hubel and Wiesel, 1959).

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Fio. 5.Records from unit activated by ipsilateral eye only; unresponsive to stationary spots, influenced by movement in an area temporal to area centralis. A slit (0-5 x 8) moved back and forth transversely with different orientations, as shown to the left. For slit orientations evoking responses only one direction was effectiveup and to the right. Time, 1 sec. (Hubel and Wiesel, 1959).

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about "a space in which any point is described in terms of how it is related to what is around it." "In a system like this, a unique combination of the four qualitative contexts in a certain spatial relation may define a class of objects. . . . Obviously, this is a way in which the universals 'prey' and 'enemy' can be recognized or inferred directly without requiring a scanning operation." These observations show how largely perceptual discrimination in vision depends upon physiological organization in the receptor organ, the retina, and this in turn upon the anatomical fact that a large number of retinal receptors converge upon a single optic nerve fibre, the ratio being according to Granit (1955) "of the order of 100:1." Only such an organization could enable a single retinal field converging on a single neural unit not only to respond to movement, but to discriminate movement in one direction from that in the opposite. Hubel and Wiesel point out some of the contributions which this peripheral discriminative organization could make to perceptual function. As Granit (1955) says, the retinal surfaces are "capable of good discrimination on the same basic principle as is the skin, i.e. by using overlapping receptive fields of different sizes, from very large to very small, the latter likely to be multiplied in cone eyes which have good visual acuity." Then there is the role of spontaneous activity in sense organs, also discussed by Granit. He says that, "to hold the view that sense organs, and in this particular connexion the retina, energize certain systems and individual cells by way of a maintained spontaneous activity is to provide a new role for peripheral inhibition in sense organs, whatever its cause. If all optic nerve fibres were silent, unless stimulated, an effect of inhibition around the edges of illuminated patches projected on to the silent retina would be negligible, because it would never be transmitted to the perceiving cortex. But if the sense organ normally is spontaneously active, the suppression of the inherent activity will be notified to the cortex. . . . The edges around the projected patches on the cortical map may now appear as regions of correspondingly altered excitability." Granit points out that "intense black is perceived only by contrast against white." It is difficult to understand how black could be perceived at all without the spontaneous activity, for, while physically it corresponds to the complete absence of visual stimuli, perceptually it is a sensation as positive as white or any colour. If colour perception corresponds, as we have every reason to believe it does, to some electrical activity in the visual cortex it is probable that the same is true of the perception of black. We have here surely an explanation of the common experience of the imperception of hemianopia. If black, the absence of light waves, were represented in the nervous system by the absence of activity in the visual cortex, destruction of an area of visual cortex should cause the missing area of the visual field

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to be seen as black, which is not commonly the case. The sensation of black must therefore depend upon a positive activity of the visual cortex. Granit also uses the vestibular responses to illustrate the value of spontaneous rhythms arising in the receptor end organs. He quotes the work of Lowenstein and Sand (1940) who demonstrated that for each canal of each labyrinth there were two opposite reactions. Rotation one way increased the spontaneous discharge, and in the opposite direction suppressed it. Granit remarks: "Consider now that there are six such canals, three in each ear, and it will be apparent that the presence of a spontaneous rhythm that can be accelerated or decelerated makes for a discrimination which, unless utilized by the brain, will be wholly meaningless. . . . The semicircular canals . . . have, as do other sense organs, only the frequency code at their disposal, grafted, however, upon a magnificent space-sensitive instrument. . . . Acceleration and deceleration of the spontaneous rhythm of discharge creates an interpretable pattern. One of our major aims as sensory physiologists is to attempt to identify peripheral 'cues' for sensory discrimination in terms of the frequency code." I could show that the same principles underlie the physiological basis of perception in all sensory modalities, but I need not do this in detail: a few illustrations will suffice. In cutaneous sensation we find one cortical unit linked with a cutaneous receptive field measuring several square centimetres (Mountcastle, 1957). As in the case of the retina, inhibitory mechanisms exist. "A cell, excited by stimulating an area of skin (the 'receptive area') may be inhibited by stimulating surrounding areas, and this influence, excited mutually in a population of cells, will result in the sharpening of the edges in the central stimulus pattern, emphasizing discontinuities and increasing contrast. . . . This kind of lateral inhibition is known in visual, auditory and somatic sensory systems" (Gordon, 1962). It is unnecessary for my present purpose to discuss the still disputed question of the specificity of sensory conduction in peripheral nerve fibres: it is generally agreed that what is important for perception is the conduction of a pattern of impulses in space and time, in which many peripheral receptors and conducting fibres take part, and which is modified by the interaction of peripheral mechanisms as well as by the receptivity of central pathways {see Melzack and Wall, 1962). Such interaction is well illustrated by Granit's dominator-modulator theory of colour vision.
SOME IMPLICATIONS OF CODING

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The essential activity of nerve fibres is the conduction of electrical impulses. They vary only in the following respects: (1) Peripherally specific receptors ensure that the nerves are excited only by particular physical stimuli. (2) The rate of conduction varies in different nerve fibres

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and appears to be related, in part at least, to their diameter. (3) The temporal pattern of sensory impulses is influenced by the intensity of the peripheral stimulus and the refractory period of the nerve fibre, and (4) peripheral afferent fibres concerned with particular modalities of sensation differ in respect of their central connexions. Thus it comes about, as we have seen, that different highly complex spatio-temporal patterns of nerve impulses are excited by individual elements in the physical object, which stimulates the nerve peripherally. So far I have not used the word "information," but we must now examine what is implied by the statement, freely used by psychophysiologists, that the pattern of nerve impulses is conveying information, and that this information is conveyed in the form of a code. Young (1962), for example, says: "If we are to understand the problem of what is stored in the memory we must examine the conditions that are necessary if the outcome of events in a nexus or channel is to provide a representation of what has gone in. . . . A representation can be so made if the events in the channel are circumscribed to certain types that have been pre-arranged to constitute what we are calling a 'code.' The making of any representation depends then upon selecting some from a specific set of physical events. These will then be recognized at the end of the channel as indicating that certain events occurred at the input." Young goes on to mention speaking and writing as examples of the employment of a code. The essence of a code, in the original sense of the word, is that it is a set of symbols which represent something else. The thing represented can be translated into the code, but when it is decoded it usually assumes its original form, the code having acted merely as a convenient vehicle for the meaning, as the morse code is used in telegraphy. But this meaning may be misleading when applied to the nervous system, in which "coding" means only the representation of information, and does not imply that the information necessarily reaches consciousness, or is decoded in the sense of regaining its original form. Technically, information is defined as "a measure of how much uncertainty has been removed by the receipt of a message" (Saltzberg, 1963). Before considering the more difficult question of human consciousness let us see how the coded information works in simpler brains. The frog is a strictly practical entomologist, its only reaction to seeing a fly being to aim its tongue at it. What the coded information derived from the retina does, therefore, is to evoke the appropriate action, and appropriate here means not only biologically suitable, but also precisely adjusted. In order that this may happen, afferent impulses must convey information about the quality and position in space of the object (the fly), and the frog must also receive information about its own position in the same space. Hence the visual information must be integrated with information about the
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position of its body derived from other sensory receptors. If, for example, the statocysts of an octopus are removed, the position of its eyes is no longer adjusted to gravity, and its trained reactions to geometrical figures are disorganized (Wells, 1960). The frog reacts to a fly because the information processed by the visual system, together with that derived from the somatic sensory system, and influenced by the animal's state of alertness, evokes the spatially appropriate motor activity. Granit makes the same point in relation to the influence of the labyrinths on eye movement. "There is no more instructive demonstration," he writes, "of the significance of pattern for discrimination. The fact that we do not discriminate vestibular impressions by perception, i.e. consciously (according to most authorities) is not a serious argument when one considers that this message is correctly interpreted in the lower centers for the eye muscles. There is no reason to regard consciousness as more than a fringe on the pattern." But even simple creatures learn by experience. How this comes about has been investigated in the octopus by Young (1962) and his collaborators. As a result of his studies of visual learning he has shown in outline how nervous impulses from the eyes may lead to movement, or the inhibition of movement, in the light of past experience, and how these activities are related to particular regions of the animal's nervous system. My object so far has been to show how complex is the part played in perception by the peripheral sensory receptors, their functional organization, and that of their conducting pathways. The physiological organization upon which sensory discrimination depends begins at the periphery with receptors individually attuned to particular physical stimuli. The grouping of these receptors in relation to afferent nervous pathways has been shown to be capable of discriminating, not only movement, but also its direction. The organization of overlapping receptive fields also plays a part in various forms of sensory discrimination, and the spontaneous activity of sensory receptors greatly enriches the scope of perceptual representation. The physiological basis of perception, then, is a code of frequency of nerve impulses, and "visual perception . . . is a dynamic act maintained by a continuous frequency variation and not by a static image on a photographic plate." "The problem of color reception," for example, "must be solved by central interpretation of information over a complex frequency code which also expresses interaction" (Granit).
NEUROLOGICAL AND PSYCHOLOGICAL TIME

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Frequency involves time, and the part played by time in the nervous system has on the whole attracted little attention. (A notable exception is the work of Gooddy, 1958, 1959.) There are two aspects of itthe use of

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time in the process of information-coding, and the coding and decoding of information about time itself. The former is the subject of collaboration between physiologists and information experts: the latter is intimately concerned with consciousness, and it is this aspect alone which I shall now discuss. Receptors and neurones are organized to respond to temporal relationships in their stimuli, and hence convey information involving time. But this does not imply that such information enters consciousness. To respond to a moving object involves information about time past and present, and even an extrapolation into the future, but neurologically all that seems necessary is the existence of receptors sensitive to the movement, and the right central connexions to evoke the appropriate motor response, as we have already seen; and the nervous system can provide this without its involving any conscious experience of time. Moreover when we describe what happens in the nervous system we are concerned with the movements of electrical impulses in space (i.e. along neurons), and though we use physical time to describe these movements, we can never extract from such an account time as we experience it psychologically. For physical space-time is a succession of instants, an eternal present which is continuously changing, and if we experienced only the present we could have no conception of time. Even a memory, when we experience it, is a present state. It is true that we distinguish it from other present states by relating it to the past, but we could not do this unless we were already in some way aware of the passage of time. This has long been familiar to philosophers and psychologists. Hughlings Jackson (1931, 1, 205) said: "Time is required for consciousness," basing this on a quotation from Herbert Spencer, and drew the conclusion that loss of consciousness in an epileptic attack is due to the rapidity of the neural discharge. "During the excessive and rapid discharge of the anatomical substrata of consciousness we could not expect that conscious states would appear; on the contrary, we should, a priori, expect loss of consciousness." Psychologists have spoken of the length of time during which consecutive events are together present to consciousness as the "specious present," a term introduced by Clay (1882). Russell (1948) devotes a chapter of his book, "Human Knowledge its Scope and Limits," to "time in experience," and Whitrow (1961) in "The Natural Philosophy of Time" discusses what is involved in the specious present or, as he prefers to call it, "the mental present." He says: "Direct perception of change, irrespective of whether it is explicitly recognized as succession, requires the simultaneous presence in our awareness of events in distinct phases of presentation. The combination of simultaneity and succession in our perception would mean that the time of our conscious experience is more like a moving line than a moving point."

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Bergson (1912) comparing physical and psychological time wrote: "There is in space neither duration (durde) nor even succession in the sense in which consciousness understands those words: each of the states described as successive in the external world exists, and thenmultiplicity has reality, only for a consciousness capable of first conserving them and then juxtaposing them by exteriorizing them in relation with one another." William James (1890) believed that brain processes set up by stimuli have a measurable duration and therefore those aroused by past and present stimuli overlap, but we are still left with the problem of how simultaneous brain states are discriminated by consciousness into past and present. This is a fundamental question for the brain-mind relationship. The mental present can be understood only against a larger background. Awareness of the passage of time seems to involve the retention of past experiences against which the ever-changing present is perceived as novelty, until it too becomes past. Thus the sense of time depends upon a qualitative difference between past and present experience. Several things follow from this. Time as a relationship of order may well be represented by a coding in the nervous system based on spatial organizationone aspect of the problem of serial order discussed by Lashley (1960). The spatial coding of time has long been a commonplace, e.g. in the gramophone record and tape recorder. In such devices spatial relations can be reconverted into temporal ones by means of movement, as no doubt happens in the nervous system. But it should be noted that past time is still coded when consciousness deals with it. When we remember past years, it does not take us years to do it: we deal with the past symbolically, whether in the form of images representing past experiences or as awareness of a relation of order between them. Here we seem to have reached a point at which we can find no physiological equivalent for what is happening in the mind. Kliiver (1962) asks "whether an 'engram of succession' (that is, of the ante- or postsignature of events) can ever be derived from a 'succession of engrams'." The answer would seem to be that awareness of succession is a mental event, but a succession of engrams is a series of physical events, which, as we have already seen, are known to be successive only by mind. An engram of succession would also be a physical event, and we cannot extract from a description of physical events something which it does not contain. In other words we cannot here at present fully translate the language of neurology into the language of psychology. Attempts have been made to do this. Dobbs (1951), discussing "the relation between the time of psychology and the time of physics," is led to postulate two dimensions of psychological time, so that events which are successive in "phase time" occur "as a whole in a single movement of transition time."

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Denbigh (1953), aware that the problem is to know how to distinguish actuality from reminiscence, "the is state from the was state," thinks that there are qualitative differences between them. But how do these arise ? From the fact he says "that we participate, in our own bodies, in the same chemical and physical processes which occur in the external world," and "the time sense of a human being derives from their existence within him, giving him the feeling of an inner time not based on the five senses." Many living organisms certainly seem to have an inherent sense of time, an inner clock by which they can regulate their activities. Though some physiological factors are known to influence this inner time its nature is not understood. What we do know about it suggests that some rhythmical activity provides a continuous flow of information which is stored to modify subsequent behaviour. But this is a physiological explanation: its translation into psychological terms still eludes us. We can hardly consider time without mentioning memory, but this is too large a subject to be dealt with adequately here; I will discuss only one or two points, which are relevant to my later discussion of consciousness. Studies of retrograde amnesia, and the effect of brain lesions on memory, suggest that both anatomically and physiologically the neurological activity underlying the mental present, and the recall of recent memories, is different from that concerned with both the long-term storage of memories, and recognition. The situation is well summarized by Adams, Collins and Victor (1962) in a recent paper. After presenting evidence that the lesion responsible for persistent retrograde and anterograde amnesia is situated in the medial thalamus they say: "The neurophysiologic mechanism for such complex types of learning as numerical relations, verbal symbols and concepts evidently is seated in the temporal lobes and upper brain-stem structures and their interconnections, in close relation to the mechanisms that maintain consciousness and attention. . . . Yet the neo-cortex itself is involved in the mechanisms of all special auditory, visual, tactile, and other learning and memories as well as those for words, mathematical figures, etc., for lesions here abolish specific memories and prevent relearning of this type of material. In other words, the rhinencephalic-diencephalic mechanism must act with parts of the cerebral cortex, as well as with the upper brain stem mechanisms of consciousness." As we learn more about this and about the role of the thalamus and hippocampus in the retention and recall of recent memories, we may expect to add to our knowledge of the brainmind relationship. * The idea of the mental present as a state enduring in time may help to bridge the gulf between brain-states and sensations. We know that an

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activity of the brain underlies our awareness of a colour, a sound, or a pain, but how, it is said, can the corresponding brain activity be a colour, a sound or a paina sensation when directly experienced, a series of nerve impulses when described by the physiologist? Consider sound: from a physical standpoint a sound is a wave-length of a certain frequency, a concept involving time. Owing to the physiological properties of the nerve fibres and the rate at which nerve impulses are conducted, it is certain that the activity of higher levels of the nervous system, for example the auditory cortex, which leads to the awareness of a sound, does not involve the reproduction of the original frequency of the sound-waves striking the ear: it is a coded representation of that frequency. The existence of the mental present seems to imply that what is presented to consciousness has a duration, and hence a structure in time, for which Whitehead's term "prehension" is a useful neutral description. A sensation may then be a prehension in time of those events in the nervous system which are presented to the neurophysiologist successively. Recent work on speech perception gives us a glimpse of this process at work. Liberman, Delattre and Cooper (1952) observed that whether a subject heard a simple unvoiced stop consonant as " p " or "k" depended in part upon the vowel by which it was followed. Hence the following vowel "plays a critical part in the auditory perception of 'p' and 'k'," and "in that event the irreducible correlate for 'p' and 'k' is the sound pattern corresponding to the consonant-vowel syllable." In this case the auditory stimuli representing the consonant and vowel are successive, and they are presented to consciousness as successive, but the quality of the earlier event, represented in consciousness by hearing the consonant, is influenced by the later one, represented in consciousness by hearing the vowel. This would be explained if the neural state set up by the first stimulus had time to be modified by that set up by the second before it entered consciousness. Thus in the mental present there is not only overlapping, but mutual modification of the representations of events, which in physical time are successive. Distortion of time sense has often been observed by subjects taking hallucinogenic drugs, especially mescaline. A number of factors may be involved, perception, inner time sense, the serial ordering of experiences, and memory. It does not seem necessary to suppose that such disordered awareness of time-relations implies awareness of other dimensions of time, as has sometimes been supposed (Mayhew, 1963). There is one other aspect of perception which, though not involving time, may conveniently be considered at this point. The sense organs and the nervous system consist of vast numbers of minute units, yet this granular structure is not reproduced in sensory experience: there is nothing

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for example in a coloured surface, or the feel of a smooth pebble, to suggest that it is mediated by a vast number of discrete neural units. Consciousness, therefore, involves spatial as well as temporal prehensions, and we can see this process at work in the activity of our sense organs, vision, for instance, converting the multiplicity of minute dots in a half-tone photographic block into gradations of light and shade. Finally, "the development of distance-receptors had a profound effect upon the organism's relationship not only to space but to time. If it is to react to objects at a distance, time must enter into the organization of its nervous system in a manner which has no parallel in the more primitive creature which responds automatically to immediate stimuli" (Brain, 1950). If activity is to be organized in time the organism needs an enduring representation of objects, and develops feelings in relation to them. A consciousness which represents objects in time must itself be based upon time, and is unsuitable for dealing with reactions which must be immediate. We cannot afford time to think before adjusting our eye movements to vestibular information: hence such reactions are unconscious.
THE PHYSIOLOGICAL BASIS OF CONSCIOUSNESS

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In the symposium, "Brain Mechanisms and Consciousness" (1954), there are many allusions to the location of consciousness. This is perhaps the most obvious question concerning its physiological basis. Where in the brain, it may be asked, is the function of consciousness to be located ? Hughlings Jackson (1931, 1, 367) said: "States of consciousness are assumed . . . to be merely concomitant with certain nervous states, those of the highest cerebral centres," but he was not very precise as to the location of these. Several facts seem to support the idea of a location of consciousness. First, lesions of some parts of the brain appear to have no effect upon consciousness, while lesions of other parts affect it profoundly. Then there are the effects of lesions of the sensory pathways. If, for example, the optic nerves are divided, the subject can no longer see, but there is no reason to suppose that any visual consciousness remains in the eyes. The same is true of the visual pathways up to the cortex, and, as we cannot trace vision beyond the visual cortex, it is sometimes said that at the cortex visual impulses "enter consciousness." Similar statements can be made about all forms of sensibility. Head (1920) in distinguishing between the sensory functions of the optic thalamus and the cerebral cortex says: "The essential organ of the thalamus is the centre of 'awareness' for certain aspects of sensation. . . . The feeling-tone of somatic or visceral sensation is a product of thalamic activity. . . . Conversely, the more entirely any aspect of sensibility depends upon discrimination the more it is the concern of the sensory cortex." Such

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statements seem to suggest that certain areas of the brain have a special relation with consciousness. Either the arrival of sensory impulses itself constitutes the corresponding sensory experience, or we are led to picture consciousness as like a person sitting at a desk supplied with a number of telephones, represented by these areas. If, however, we examine this rather naive picture, we see that for several reasons it cannot represent the true state of affairs. First, is there any reason to distinguish the cortical end-station of a sensory pathway from any other point of the pathway leading up to it? Though it is possible to distinguish differences in the disorder of function related to the level of the lesion, inherently the loss of function is the same, and, as has been shown above, we cannot now maintain that discriminative activity is solely a function of the sensory cortex. "The visual cortex is a necessary condition of vision, for its destruction renders the subject blind, but the same is true of the optic nerves. Why, then, do we attach importance to the cortex? Because it is the last anatomical point on the afferent visual pathway of which this is true. But it does not follow that the function of vision is locatedthere. It may be inaccurate to speak of its location in any particular part of the nervous system. If the brain largely acts as a whole, vision may be a property of the whole of it, in which case it is represented in the brain as a whole in a different way from its representation in the visual cortex. The latter then becomes only the last point at which nerve-fibres solely concerned with vision exist as an isolable group. The same may be true of other sensory pathways, and of the motor cortex, and motor pathways too" (Brain, 1960). In that case we should not expect localized lesions of the brain outside the visual cortex to affect vision in the same way as lesions of the visual cortex, but we might expect lesions of other parts to influence visual reactions as, in fact, they do. But there is another form of naivete" about the localization of consciousness which arises not unnaturally from the facts of anatomy and physiology. The physiologist knows that sensory impulses initiated at the retina, or at some particular segment of the body, cause electrical impulses to be transmitted to an area of the cerebral cortex in which that particular part of the retina, or of the body, is said to be represented, and the clinician observes a similar correlation between destruction of these areas and the localization of the loss of function which results. But it does not follow that the arrival of sensory impulses at the cortical sensory area is all that is required for them to enter consciousness. "If we imagine the nervous pathways from the toe to the cortex completely isolated from the rest of the nervous system, it is clear that they could by themselves convey no localizing information, for how could electrical impulses exactly like those going to all other cortical sensory areas mean that something is happening to the toe? This surely provides a clue to

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the answer, which is that we can localize touch only by perceiving it in relation to a representation of the body as a wholewhether in the form of unconscious schema or fully conscious image is unimportant for our present purposeand this body-schema maps out, as it were in a set of co-ordinates, all the possible voluntary actions which we should need to take in order to remove the stimulus either with the other foot or with either hand or in any other way. . . . This process of localization of touch is a cortical function, but if I am right in thinking that it relates the touch felt to the body-image as a whole, it must involve pathways far more extensive than the primary sensory areas for touch in the cortex. When, therefore, the cortical area corresponding to one great toe is excited by the arrival there of afferent impulses resulting from stimulation of the toe itself, such excitation must cause a widespread irradiation of impulses linking the toe area with the parieto-occipital areas which are concerned with awareness of the body-image, not only in the same cerebral hemisphere but by way of the corpus callosum with the corresponding area on the opposite side of the brain " (Brain, 1950). This, no doubt, is what Hughlings Jackson (1931, 1, 350) meant when he said: "Just as any state of consciousness is a state of a whole person psychical, so the correlative activity is of nervous arrangements representing a whole person physical (the whole organism)." To avoid possible misunderstanding I must add at this point that the statement that "the brain acts largely as a whole," does not conflict with the evidence for a high degree of specialization of function in certain areas. Apart from the special sensory and motor areas, and areas concerned with speech, spatial awareness, etc., there is abundant evidence that some regions, e.g. the ascending reticular formation, are particularly concerned with the function of consciousness as such, while other areas, e.g. the neocortex, are concerned primarily with its content: moreover, the damaged brain may continue to function and provide consciousness, and a content for consciousness, in however limited a manner. Nevertheless there is also evidence that large areas of the cortex show a response to a single sensory stimulus (Jasper, Ricci and Doane, 1960), as anticipated by Lashley (1960), and may well play a part in memory storage, so that in normal circumstances the nervous system does act largely as a whole.
THE NATURE OF CONSCIOUSNESS

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Kube (1954) said: "Consciousness of anything implies and depends on an ability to differentiate an T from a 'non-I' world." This is fundamentally true, but it is only half the truth, the other half being that consciousness of anything implies, and depends on, a fusion of the subjective and the objective.
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Consider the essential elements in the simple experience of seeing a table. Neurologically, in order that this may happen it is necessary that the image of the table falh'ng upon the retinas should arouse nerve impulses which eventually lead to changes in the neurons of the visual cortex. Much more than that, of course, is required, but for our present purpose we can neglect the rest. From one point of view all this is subjective it is happening in the brain of the subject, and, correspondingly, the conscious process of seeing a table is subjective: it is my seeing. But it is also objective, because it includes structural elements which it shares with the table, and by means of which it conveys information about it. Without the preliminary fusion of the subjective and objective in consciousness there could be no differentiation of an " I " from a "non-JL" world. In this process of differentiation consciousness may label, as objective, sensory qualities, such as colours, which are solely products of the activity of the nervous system. Hence the distinction between what is subjective and objective in experience may not correspond to the distinction between events occurring outside the subject's body and those taking place in his nervous system. Yet one fundamental distinction between the " 1 " and the "non-1" worlds is determined by the boundary of the human body. All action in the external world requires and implies information as to the existing orientation of the body in relation to it, and of the parts of the body to one another. Such information may be related to consciousness in three ways. It may be permanently unconscious, as in the case, for example, of afferent pathways to the cerebellum. It may be largely unconscious, but potentially conscious, as we may use our legs in walking while thinking about something eke, and yet can direct our attention to the position of our feet and so become conscious of them. The third way is that by which the information contributes to a totality of which we are conscious, though we are not aware of its specific contribution, as our labyrinths contribute to our perception of our own posture, and our awareness of the external world. We cannot normally detect this contribution, but, if it goes wrong and causes an attack of vertigo, or if both labyrinths are destroyed, we soon become aware of the importance of the information which has been lost. But what of self-consciousness? What of our consciousness of thoughts, feelings and memories? These come and go. In so far as they are my own they are part of myself, but it may be argued there must be a self which is in some way independent of particular thoughts, feelings and memories, and that being conscious of those involves experiencing them in relation to this fundamental self. The analysis of consciousness by the disentangling of these factors calls for much more psychophysiological and pathological observation; and the study of the effects of drugs

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which alter perception, self-awareness, memory and the consciousness of time may be especially fruitful.
CONCLUSIONS

Several conclusions can be drawn from the facts we have just been considering. (1) First, neurologically, consciousness would seem to involve an interrelationship between the neural basis of the body-schema, and the perceptual organization by which the external world is represented. (2) Equally important are the processes by which experiences become recorded in memory, and the neural basis of the emotions; and thought and speech may also be involved to a variable extent. (3) The everchanging content of consciousness means that the extent of the brain in action, particularly perhaps of the cerebral cortex, must be equally variable. (4) If there are nodal areas, they must surely be diencephalic, since it is in the brain-stem and diencephalon that the neural bases of attention, emotion and memory seem most closely related to one another. (5) But we have long known that functions are not to be located exclusively where lesions disturb them, and cortex and diencephalon need to be regarded "not as a hierarchy, but as an integrative unity, such as that represented by the yin and yang of Taoism intertwined in their selfembracing circle" (Brain, 1958). (6) Finally, since consciousness takes time, we must surely picture it as a function of the sustained activity of vast numbers of neurons extending over large areas of the brain. Hughlings Jackson (1931, 2, 85) said: "I take consciousness and mind to be synonymous terms. . . . If all consciousness is lost mind is lost. Unconscious states of mind are sometimes spoken of, which seems to me to involve a contradiction. That there may be activities of lower nervous arrangements of the highest centres which have no attendant psychical states, and which yet lead to next activities of the very highest nervous arrangements of those centres whose activities have attendant psychical states, I can easily understand. But these prior activities are states of the nervous system, not any sort of states of mind." What I set out to do was to show how much the process of perception depends upon the activity of peripheral receptors and conducting pathways. Quotations have shown that neurophysiologists working in this field find it difficult to describe their results without using psychological terms. As Granit (1955) says, "on the whole . . . the distinction between sensation and perception has lost its validity. . . . A sensation is an exceedingly complex affair." We have seen, too, how hard it is to draw the line between conscious and unconscious factors even in perception, and we are familiar with our own dependence upon unconscious processes for our comprehension and utterance of speech, and for thought.

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Jackson's definition of mind in terms of psychophysical parallelism led him to say that unconscious mind is a contradiction. Sherrington (1940), an interactionist, was so impressed with the kind of facts I have been describing that he said that "there would seem to be a grade or grades of mind which we do not experience, as well as the mind which is our mental experience." These questions involve "the logical analysis of the conventions of language," which I shall not now discuss. But the new physiological facts we have been consideringand many others surely encourage us to use the terms mental and physical interrelatedly at progressively lower levels of nervous function. I began by talking about the contribution of peripheral sensory pathways to perception: let me end with a quotation which puts into a focus of relationship much of what I have said. "In the genesis of visual perception," writes Sperry (1962), "problems of chemical coding are involved in the differentiation of retinal specificity, the chemotactic guidance of the optic axons to their central terminals, their selective synapsis with central neurons, and the selective synapsis of these in turn with deeper elements of the system. The chemical tagging of the neurons for perception of directionality, for color discrimination, for brightness, and for the on-off and other physiological properties requires several dimensions of chemical specificity within a single neuron. . . . It is quite possible that the chemical changes associated with learning and memory in the mammalian cerebrum represent an evolvement, more or less direct, of these basic properties underlying selective chemical affinity." In these sentences Sperry shows us genetic information-coding preparing the way for perceptual information-coding, learning and memory, and suggests that they are parts of the same language. The language Sperry is using is a physical one, but he cannot describe what the physical changes are doing without using the psychological terms, perception, learning and memory. Hughlings Jackson was not an original philosopherhe took over a philosophy ready-madebut we commemorate him as physician, physiologist and Dsvcholoeist. and surely we cannot do that better than by continuing to study those correlations between mental and bodily activity which he found so absorbing. Only so, I believe, shall we reach a better understanding of the nature of their relationship. I am grateful to Professor Ragnar Granit and Drs. Hubel and Wiesel for permission to reproduce illustrations from their publications.

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SOME REFLECTIONS ON BRAIN AND MIND REFERENCES

401

ADAMS, R. D., COLLINS, G. H., and VICTOR, M. (1962) In "Physiologie de l'Hippo-

campe." p. 273.

Editions du Centre National de la Recherche Scientifique.

Paris,

BERGSON, H. (1912) "Essai sur les Donnees Immediates de la Conscience."

1 lth Edn.

Paris, p. 91.
BRAIN, W. R. (1950) Brain, 73, 465.

(1958) Brain, 81, 426.

(1960) Brit. J. Phil. Sci., 11, 177. CLAY, E. R. (1882) "The Alternative. A Study in Psychology." London, pp. 167-8. Council for International Organizations of Medical Sciences (1954) "Brain Mechanisms and Consciousness." A Symposium, Ed. by J. F. Delafresnaye, Oxford, 1954. DENBIGH, K. G. (1953) Brit. J. Phil. Sci., 4, 183. DOBBS, H. A. C. (1951) Brit. J. Phil. Sci., 2, 122 and 177. GOODDY, W. (1958) Lancet, 1, 1139. (1959) Lancet, 2, 1155. GORDON, G. (1962) In "The Assessment of Pain in Man and Animals." Ed. by C. A. Keele and R. Smith. Edinburgh and London, p. 123. GRANIT, R. (1955) "Receptors and Sensory Perception." New Haven and London" HEAD, H. (1920) "Studies in Neurology." London.
HUBEL, D. H. (1959)/. Physiol., 147, 226.

Downloaded from http://brain.oxfordjournals.org/ at Bodleian Library on February 11, 2013

(1963) /. opt. Soc. Amer., 53, 58. D. H., and WIESEL, T. N. (1959) /. Physiol., 148, 574. , (1960)/. Physiol., 154, 572. , (1961) / . Physiol., 155, 385. , (1962) / . Physiol., 160, 106. , (1963) /. Physiol., 165, 559. JACKSON, J. H. (1931) "Selected Writings." Ed. by J. Taylor, London, vol. 1. JAMES, W. (1890) "The Principles of Psychology." New York, vol. 1, p. 635. JASPER, H., RICCI, G., and DOANE, C. (1960). "Moscow Colloquium on Electroencephalography of Higher Nervous Activity." Ed. by H. H. Jasper and G. D. Smirnov, Electroenceph. clin. Neurophysiol. Suppt. 13, p. 137. KLUVER, H. (1962) In "Macromolecular Specificity and Biological Memory." Ed. by F. O. Schmitt, Cambridge, Massachusetts, p. 94. KUBE, L. S. (1954) In "Brain Mechanisms and Consciousness." Ed. by J. F. Delafresnaye, Oxford, p. 444. LASHLEY, K. S. (1960) In "The Neuropsychology of Lashley." Ed. by F. A. Beach, D. O. Hebb, C. T. Morgan and H. W. Nissen. New York. LIBERMAN, A. M., DELATTRE, P., and COOPER, F. S. (1952) Amer. J. Psycho/., 65, 497.
HUBEL,

LOWENSTEIN, O., and SAND, A. (1940) / . Physiol., 99, 89.

MATURANA,

H. R., LETTVIN, J. Y., MCCULLOCH, W. S., and PITTS, W. H. (1960) /. Gen. Physiol., 43, July Supp. p. 129.

402
MAYHEW,

LORD BRAIN

C. (1963) In " Hallucinogenic Drugs and Their Psychotherapeutic Use." Ed. by R. Crocket, R. A. Sandison and A. Walk. London, pp. 170-173. MELZACK, R., and WALL, P. D. (1962) Brain, 85, 331. MOUNTCASTLE, V. B. (1957) /. Neurophysiol., 20, 408. RIDDOCH, G. (1917) Brain, 40, 15. RUSSELL, B. (1948) "Human Knowledge: Its Scope and Limits." London, p. 226. SALTZBERG, B. (1963) In "Information Storage and Neural Control." Ed. by W. S. Fields and W. Abbott. Springfield, Illinois, p. 5. SHERRINGTON, C. S. (1940) "Man on His Nature." Cambridge, p. 306. SPERRY, R. W. (1962) In "Macromolecular Specificity and Biological Memory." Ed. by F. O. Schmitt. Cambridge, Massachusetts, p. 70. WELLS, M. J. (1960) /. exp. Biol., 37, 489. WHITROW, G. J. (1961) "The Natural Philosophy of Time." London. YOUNG, J. Z. (1962) J. ment. Sci., 108, 119.

Downloaded from http://brain.oxfordjournals.org/ at Bodleian Library on February 11, 2013