The Neuropsychology of Consciousness

England

PREFACE

In September 1990, with the support and encouragement of the Russell Trust, and with additional financial help from the Wellcome Trust, a symposium entitled ‘Consciousness and Cognition: Neuropsychological Perspectives’ was held at the University of St Andrews. The intention was to assemble a group of the major researchers at the forefront of this field. The meeting proved a great success, and we hope that the chapters of the present volume will help to explain why by conveying some of the excitement of current research and theory in the area.

The starting point for the symposium and for the book was the widespread realization that in several areas of human cognition (e.g. visual perception, memory, language comprehension, and attention), the severe and profound impairments due to brain damage that have been described over the past 150 years are often not absolute. In particular, the use of indirect methods of testing may reveal unsuspected preservation of capacities that are undetected by more traditional direct methods. Since such techniques reveal abilities of which the patient is unaware, these findings have sometimes proved as much a surprise for the patient as for the investigators, and have thrown questions about the relationship between cognition and consciousness into sharp relief. Lawrence Weiskrantz was a major pioneer in what are still the two most intensively investigated of the neurological deficits that give rise to dissociations between ‘aware’ and ‘unaware’ processing: amnesia, and ‘cortical blindness’. Furthermore, his contributions since those early reports have continued to be highly influential. It is therefore particularly appropriate that he should provide the Introduction to the present volume.

We would like to express our thanks not only to the sponsors of the symposium, and to those contributors to it who are represented in this volume, but also to others whose contributions at the meeting undoubtedly, if covertly, helped shape the contents of the book. We particularly acknowledge the contributions of Daniel Bub, Andrew Ellis, Melvyn Goodale, Michael Kopelman and Patrick Rabbitt. Finally, we thank Andrew Carrick of Academic Press for his organizational help, and Nikki Jones for her untiring secretarial assistance; without them this book could not have been completed in so short a time.

A.D. MILNER and M.D. RUGG,     St Andrews

April 1991

CHAPTER 1

Introduction: Dissociated Issues

Lawrence Weiskrantz

Publisher Summary

This chapter discusses various issues related to dissociations. Capacity in the absence of, or independent of, explicit awareness has come under heavy scrutiny in phenomena such as repetition priming, backward central masking, and lexical decision. Evidence for distinctions between residual capacity and acknowledged awareness is often based on single dissociations, while independent systems or processes require the use of double dissociations. Therefore, especially in the area of subliminal perception, quantitative differences can be reliably measured between objective discriminative thresholds and the threshold for subjective awareness. Such investigations raise the question of whether it is possible to measure awareness or nonawareness differences in animals. Each of the neuropsychological domains within which awareness or nonawareness dissociations have been reported has generated domain-specific dissociative descriptions.

1.1

IMAGING AND EPIDEMIOLOGY

Every period in the relatively brief history of neuroscience has considered that it had a special, sometimes even ultimate, insight into the neural mechanisms of conscious awareness, offered against a sustained background hum of denials that the problem was real, or, if real, capable of being investigated within science. In the main, among those who accepted it as both real and of scientific interest, the approaches were largely large-scale correlational, e.g. changes in electrical activity during periods of heightened attention, or during stages of sleep, as in the reticular formation research, combined of course with supportive clinical and behavioural invasive research. Alternatively, they were theoretical and speculative, often depending upon an implicit assumption that awareness depends upon particular details of cortical activity, an assumption rooted in late nineteenth century thinking but extending into modern times.

While modern work is continuous with its past, the last 15–20 years has nevertheless seen two developments which make the period into which we are about to enter somewhat special and exciting, and a volume such as this one especially useful and topical. The first of these is the development of imaging techniques which, even now, strike one as so remarkable as to exude a flavour of science fiction. Combined with increasing sophistication of analysis of event-related potentials, they provide a heretofore totally inaccessible view of fine-grained patterns of brain activity that occur during ongoing cognitive activity. Even more significant for the present theme, and even more recent, they are being put to the service of those asking sophisticated experimental questions about the information processing aspects of those cognitive operations, so well illustrated in the work of Michael Posner (see Chapter 5, this volume).

A second development, one that is unique to this age, is the virtual epidemic of dissociations discovered by neuropsychologists whereby residual processing occurs in the absence of acknowledged awareness. The first to be recognized, perhaps, was in the field of amnesia, where it is now widely agreed that amnesic patients show good retention for certain types of events even though the patients acknowledge no memory for them as such. But other examples have recently come to light, most of them also the subject of detailed treatment in this volume: blindsight, blind-touch, ‘deaf hearing’ (Michel, 1990), prosopagnosia and other forms of agnosia, dyslexia, unilateral neglect, and aphasia. A complementary condition, the denial of loss of a capacity (anosognosia), has also come under focused investigation. Potentially it seems possible that the dissociation between capacity and acknowledged awareness will emerge for every cognitive, and even non-cognitive, neuropsychological syndrome.

This epidemic of dissociations encouragingly has led to an increase in interest by philosophers of mind in empirical evidence; without experimental analysis none of the dissociates would be known. In addition, they have also interacted with, indeed often been the spur for uncovering, apparently similar evidence in the human experimental psychology of normal subjects. Capacity in the absence of, or independent of, explicit awareness has come under heavy scrutiny in such phenomena as repetition priming, backward central masking, lexical decision, stem-completion, picture completion, sentence arrangement, anomalous pictures/sentences, long-delayed forced-choice recognition, subliminal perception, skills, autonomic responses, and shadowing in dichotic listening.

Of course, there is a wealth of detail to consider and uncover, but in the context of a volume such as this a number of more general questions arise, and two of them arise straight away. The first is how well do the phenomena within either neuropsychology or within normal human experimental psychology map onto each other? A second is how well do these two domains map onto each other? An epidemic does not necessarily imply common mechanisms. How readily can one assume a parasitism of explanation for one category of effects on another; are there common generalizations?

1.2 LOGICAL POWERS AND LIMITATIONS OF DISSOCIATIONS

All of the neuropsychological evidence mentioned above depends on dissociations, i.e. different capacities being differently affected by different treatments. Further questions emerge from a consideration of the logical powers and limitations of dissociations. Evidence for distinctions between residual capacity and acknowledged awareness are often based on single dissociations, e.g. it is readily possible to demonstrate a loss of ‘explicit’ recognition without a loss of priming, but apparently not vice versa. Single dissociations are useful logically for drawing inferences about hierarchical control, but provide a very weak basis for arguments in favour of independent systems or processes, for which double dissociations are necessary (but not sufficient). Yet theories of independent systems or independent processes are not uncommon across the range of phenomena considered above, and have led to the drawing of a number of dichotomous contrasts. Which of the phenomena would we expect to be susceptible to double dissociations and for which would double dissociations be unexpected or even impossible?

Where double dissociations are found, as is the case in sub-species of blindsight, for example, inferences about independent visual processors or ‘modules’ readily emerge and are conveniently mapped onto physiological and anatomical evidence. However, even when such potentially independent processes are inferred, evidence of double dissociations tells one little or nothing about the interactions of such systems or processes. For this one needs a different type of enterprise, with theoretical underpinning.

Finally, many of the double dissociations inevitably are ‘trend’ (Shallice, 1988) or ‘relative’ (Weiskrantz, 1989) dissociations, i.e. not absolute or pure. Inferences of independence are still possible, but require qualifications based on, say, intermingling of anatomical pathways; but it is even possible that such ‘trend’ dissociations could be found in a ‘lesioned’ PDP network (see Shallice) where independent systems are, by definition, not available.

1.3 QUANTIFIED VALIDATION OF AWARENESS–NONAWARENESS DIFFERENCES

It is now becoming recognized, especially in the area of subliminal perception, that quantitative differences can be reliably measured between objective discriminative thresholds and the threshold for ‘subjective awareness’. Moreover, Cheesman and Merikle (1986), having measured such a difference, also validated the difference in terms of converging evidence from differential stimulus control of another phenomenon, the effect of varying probability of congruent information in a Stroop-priming task. More recently, Kunimoto et al. unpublished data have subjected the difference between objective and subjective thresholds to a signal detection analysis by requiring a retrospective judgment of correctness after each trial. Their results yield a somewhat different outcome from Cheesman and Merikle, but they nevertheless found a small, but highly reliable, subliminal effect in colour identification by normal subjects.

Given this background, a number of questions arise in relation to both the neuropsychological evidence and that from human experimental psychology. How many of the phenomena from those domains could be quantified in this way? When quantified, could the difference in thresholds be shown to be qualitatively dissociable, i.e. how much converging evidence has been generated, or even could be generated? Is such a qualitative functional difference necessary? A further question arises in some neuropsychological domains: how can one deal quantitatively with the phenomenon of ‘switched’ or ‘gut’ awareness? For example GY, a blindsight subject, sometimes says (but this is not true of all blindsight nor of GY under all conditions) that he ‘knows’ that a stimulus has occurred, but insists that he definitely does not ‘see’.

A final and deep question concerns whether it is possible to measure awareness/non-awareness differences in animals. The question is seen to be real when it is realized that some of the methods used to establish such a difference in human subjects can be mapped directly on ‘gedanken’ experiments with animals. For example, the method used by Kunimoto et al. (unpublished data) is logically closely similar to a suggestion for independent response alternatives for forced-choice discriminations, on the other hand, versus responses on each trial for ‘seeing’ and ‘guessing’, on the other, that could be used with monkeys (Weiskrantz, 1986, 1988; see also Cowey & Stoerig, Chapter 2, this volume).

1.4 DESCRIPTIVE CHARACTERIZATIONS

1.4.1 Dissociative Descriptions

Each of the neuropsychological domains within which awareness/non-awareness dissociations have been reported has generated domain-specific dissociative descriptions, e.g. in the memory field, implicit versus explicit, procedural versus declarative, semantic versus episodic; in vision, two-visual system distinctions between detection and identification, ‘where’ or ‘whether’ versus ‘what’; in prosopagnosia, ‘directed visual processing’ versus ‘face recognition’, and in aphasia, ‘on-line’ versus ‘off-line’ processing.

Do these various dichotomous pairs have anything in common, other than that one in each is said to lack acknowledged awareness? Do we proceed among the set for analysis one-by-one, or are bolder generalizations possible? Given that some of the dichotomies emerge from single dissociations, and others from double dissociations, there is a primafacie case for separate examination and, at the very least, a dichotomous classification of different dichotomies. At a more abstract level, a central but difficult question is whether one can distinguish between disconnected awareness and subtraction of a particular sub-system. For example, in amnesia can we account for residual function by subtraction of a system specialized for, say, episodic memory, or are we dealing with a disconnection from a central ‘awareness’ module? At this stage, there are no clear, sharp criteria for deciding between these two alternatives.

1.4.2 Graded Processes

In contrast to dissociative descriptions is an approach in terms of graded processes. Depth of processing, data-driven versus concept-driven processing, and transfer of procedures are all recent examples of attempts to explain the apparent disjunctions in human experimental psychology and neuropsychology in terms of continuously variable aspects of task demands and subjects’ capacities (cf. Roediger et al., 1989). A more traditional example in another domain is levels of vigilance as a concept in attention.

A question that naturally arises is whether there is any way to choose between dissociative and graded accounts. It is agreed that double dissociations are necessary but not sufficient to prove that independent and disjunctive processes or systems exist, and so what else is needed? It is also the case that, whether or not one’s allegiance is to multiple systems or to graded processes, all relevant research starts from results using tasks that are impure for either type of category: there is no such creature as a ‘pure task’, and hence further dissection beyond a catalogue of tasks, as such, is mandatory (cf. Weiskrantz, 1989; Jacoby & Kelley, Chapter 10 this volume). Can we get beyond a show of hands? Given that the decision about independent systems or processes will be based, in part, on convergent evidence, what kind of pragmatic and convergent evidence is available? In neuropsychology we at least have independent evidence from neuroscience; e.g. for claims of independent varieties of blindsight, one can bolster one’s case by reference to independent anatomical outputs from V1 to V2–V5, together with inputs from the superior colliculus and the lateral geniculate nucleus to V2–V5 that bypass V1, but what kinds of converging evidence within the domain of normal human experimental psychology are available to which a comparable appeal can be made?

1.5 MULTI-DOMAIN CHARACTERIZATIONS

Some theoretical approaches have assumed that there is a unified nature of ‘conscious awareness’ that warrants a general unified account in neurological or physical terms. Many dissent from such a position, on various grounds: philosophical, semantic, pragmatic, over-complexity and inhomogeneity even in folk-psychology discourse. Among those that find the question of the nature of conscious awareness viable, there are suggestions at widely different levels and different degrees of specification. A traditional view, given impetus by Sperry (1969), is that conscious awareness emerges directly as a function of complexity of the nervous system, but without specifying what it is about complexity that allows such a qualitative attribute to arise. Other views have tried to specify particular hypothetical arrangements within nervous systems, or computational systems, independent of whether some threshold level of complexity as such is a necessary condition. Johnson-Laird’s (1988) view of parallel hierarchical systems might be an example of this. Another view, similarly placed within a hierarchical arrangement, characterizes systems with ‘awareness’ as those with parallel monitoring circuits which, in turn, also have communicative links, especially with language, and have an effect on control systems (Weiskrantz, 1988). This in turn is consonant with philosophical views of awareness involving ‘higher order thoughts’, as advanced by Rosenthal (1986).

An account that was deliberately designed to assimilate neuropsychological evidence of dissociations is the DICE (‘dissociable interactions and conscious experience’) schema of Schacter (1989), a box-and-arrow model which assumes the existence of a conscious awareness system with specific connections to knowledge modules and an episodic memory system (among others). It is a moot point whether the conscious awareness system as such is necessary or whether multiple sub-systems will themselves be sufficient. But even if one assumes the latter, questions will continue to nag us as to what it is about particular sub-systems that enables one to attach an attribute of phenomenal awareness to their operation. Can one point to any stage or organizational property of such systems and say that is where, or how, the critical step takes place–or is this a pseudo-problem?

A quite different approach is that of Crick and Koch (1990) who link the question of awareness to the ‘binding’ problem, the problem of how the multiplicity of specialized visual neurons allows a single coherent visual image to emerge. Attentional mechanisms that selectively focus on different details of, say, visual inputs to V1 are assumed to be reflected in interactions among different widely dispersed neurons throughout the whole galaxy of specialized visual cortical areas. It is a theory that links awareness to selective attention and, it is assumed, also to short-term, ‘working memory’. It may be that particular lesions will disenable specific forms of binding to take place, and hence lead to specific forms of ‘unawareness’. It is a moot point whether patients with attenuated ‘working memories’ have attenuated consciousness, and also whether selective attention may or may not still operate in unaware vision in blindsight or in on-line semantic processing in aphasia or prosopagnosia, but the schema does lead to some clear testable predictions.

It is worth bearing in mind, in case we were allowed to forget them, some current popular accounts that take us into quite different ballparks, especially those based on quantum effects (Penrose, 1989) which, in turn, can lead to various species of interactionistic dualism (Eccles, 1987). These all assume that there is a problem of consciousness to be explained, but take it well outside any explanatory level that would link specific neural circuits with specific dissociations. Indeed, given the universality of quantum effects (actually given empirical underpinning in the limiting sensitivity of the retina), the question arises as to whether there are any parts of the nervous system (including the retina) that are not conscious.

Aside from the ‘other ballparks’ accounts of the last paragraph, the question is whether any of these approaches are any more than heuristic at the moment. All of them stimulate certain lines of enquiry and, in limited domains, testable predictions, but most are still at the level of ‘points of view’. A further question that requires consideration is the relation of any of the accounts to executive control. In lay language, what is the relation of awareness to the question of determinants of action, e.g. ‘free’ or ‘not free’ will.

1.6 SUGGESTED NEURAL CORRELATES AND EMBODIMENTS

No one would claim that there are more than hints about embodiments at the moment, although there is an air of anticipation that we are on the verge. But among the various references to the r.n.s. (the real nervous system) one can note the prominence of the parietal lobe and frontal lobe – especially its medial aspect, the anterior cingulate – in connection with neglect and attention. There is much consideration, also, of the role of the midbrain, especially the superior colliculus, in mediating certain ‘implicit’ aspects of perception, as in blindsight, although the midbrain must be considered, for this example, in relation to specialized cortical areas beyond V1 to which the midbrain (but also the lateral geniculate nucleus) projects. This, in turn, leads to speculations that ‘awareness’ may require a convergence of both the midbrain and cortical inputs.

The approach through the ‘binding’ problem and selective serial attention of Crick and Koch (1990) draws heavily on observations of coherent, correlated oscillations, roughly at 40 Hz in the firing patterns of cells in the visual cortex. Finally, mention should be made of the various studies of Libet (1982) and his colleagues of the claim that awareness arises slowly in sensory systems, as compared with the control that such systems might exert on motor responses. It is not clear just what it is about the minimal ‘on-time’ that is necessary for awareness to arise, and just what happens during this period, but Libet is one of the few physiologists making an effort to investigate temporal aspects of conscious versus unconscious processing. In addition, of course, one must mention the large literature on average evoked potentials that are related to different aspects of cognitive processing, within which domain it ought to be possible in principle to separate overt from covert modes of processing.

1.7 WHAT IS IT FOR?

Discussion of the possible evolutionary value of conscious awareness has been continuous for over 100 years, at least among those that assume that it is a special attribute that both can evolve and can bestow benefits. One dominant theme is to attach its benefits to the benefits of active thought itself – in allowing the initiation of predictive strategies, in detaching the observer from an immediate dependence on current inputs, by allowing current inputs to be linked, with or without imagery, to other events distant either in space or time, and in allowing flexible rather than automatic processing. Certainly the neuropsychological patients who suffer from any of the syndromes that are discussed in this volume are deeply disabled: amnesic patients cannot survive on priming or conditioning alone, nor can prosopagnosic patients overcome their acute embarrassment in not recognizing their children, even though they may be able to process ‘implicitly’ their semantic status, nor can the blindsight subject avoid bumping into lamp-posts, even if they can guess their presence or absence in a forced-choice situation, nor can they compare their ‘covert’ detection with past visual experience. All of these subjects lack the ability to think about or to image the objects that they can respond to in another mode, or to inter-relate them in space and in time; and this deficiency can be crippling.

A sub-species of the evolutionary position considers the very special benefits in the social domain of being able to entertain a theory of ‘other minds’, i.e. to make predictions based on what the cognition of another being is inferred to be. This, in turn, leads one to consider the disadvantages, as well as the advantages, of conscious awareness, and to the view that paranoia is a uniquely species-specific attribute: only in the case of humans is existence potentially deeply distorted by attribution of devious motives to other beings. It is not clear, from this point of view, that when one considers the collective paranoia exhibited in certain unstable political arenas, the advantages always outweigh the disadvantages.

Finally, having drifted rather far from the main theme of the book, it is time to return: it is clear that the combination of rapidly developing imaging techniques plus the extraordinary epidemic of dissociations between ‘overt’ and ‘covert’ processes, both in neuropsychology as well as in the human experimental laboratory, will generate a new and important set of insights into the mechanisms underlying forms of awareness, both functionally speaking and in terms of neural events and circuits themselves. It is encouraging to witness not only the emergence of fresh directions within experimental psychology, neuropsychology, and their partners in neuroscience in a collective enterprise, with a sophisticated dissection of evidence, but also to provide philosophers of mind with material that by its very nature will remove them from a complete dependence upon the armchair. It will be for the future to see just how dissection and assimilation in the larger arena takes place.

REFERENCES

Cheesman, J., Merikle, P. M. Distinguishing conscious from unconscious perceptual processes. Canadian Journal of Psychology. 1986; 40:343–367.

Crick, F., Koch, C. Towards a neurobiological theory of consciousness. Seminars in the Neurosciences. 1990; 2:263–275.

Eccles, J. Brain and mind, two or one? In: Blakemore C., Greenfield S., eds. Mindwaves. Oxford: Basil Blackwell, 1987.

Johnson-Laird, P. N. A computational analysis of consciousness. In: Marcel A.J., Bisiach E., eds. Consciousness in Contemporary Science. Oxford: Oxford University Press, 1988.

Kunimoto, C., Miller, J. & Pashler, H. Perception without awareness confirmed: a bias-free procedure for determining awareness thresholds. Department of Psychology, University of California, San Diego (unpublished manuscript).

Libet, B. Brain stimulation in the study of neural functions for conscious sensory experiences. Human Neurobiology. 1982; 1:235–242.

Michel, F. (1990). Hemi-anacusia is usually unknown to the patient. Poster presented at Russell Trust symposium, St Andrews, Scotland, September 1990.

Penrose, R.The Emperor’s New Mind. Oxford: Oxford University Press, 1989.

Roediger, H. L., III., Weldon, M. S., Challis, B. H. Explaining dissociations between implicit and explicit measures of retention: a processing account. In: Roediger H.L., III., Craik F.I.M., eds. Varieties of Memory and Consciousness: Essays in Honour of Endel Tulving. Hillsdale, NJ: Erlbaum, 1989.

Rosenthal, D. M. Two concepts of consciousness. Philosophical Studies. 1986; 49:329–359.

Schacter, D. L. On the relation between memory and consciousness: dissociable interactions and conscious experience. In: Roediger H.L., III., Craik F.I.M., eds. Varieties of Memory and Consciousness: Essays in Honour of Endel Tulving. Hillsdale, NJ: Erlbaum, 1989.

Shallice, T.From Neuropsychology to Mental Structure. Cambridge: Cambridge University Press, 1988.

Sperry, R. A modified concept of consciousness. Psychological Review. 1969; 76:532–536.

Weiskrantz, L.Blindsight. A Case Study and Implications. Oxford: Oxford University Press, 1986.

Weiskrantz, L. Some contributions of neuropsychology of vision and memory to the problem of consciousness. In: Marcel A.J., Bisiach E., eds. Consciousness in Contemporary Science. Oxford: Oxford University Press, 1988.

Weiskrantz, L. Remembering dissociations. In: Roediger H.L., III., Craik F.I.M., eds. Varieties of Memory and Consciousness: Essays in Honour of Endel Tulving. Hillsdale, NJ: Erlbaum, 1989.

CHAPTER 2

Reflections on Blindsight

Alan Cowey and Petra Stoerig

Publisher Summary

This chapter discusses the phenomenon of blindsight. This phenomenon has attracted the attention of philosophers, psychologists, and neuroscientists because it highlights the nature of covert vision, indicates that striate cortex is indispensible for visual awareness, and provides a means of studying the visual information carried by pathways other than the major route through striate cortex. Denial of visual experience may be present even when the subject’s detection or discrimination approaches 100% correct, when localization of an unseen target is excellent, or when the threshold for detection is reduced by less than a log unit. It has been seen that the presentation of a line stimulus in the blind field influences judgments about the relative distance of targets presented in the normal field and that the color of targets in the seeing field is affected by unseen colors in the scotoma. Some patients report occasional sensations, which may even be of a visual nature. The chapter presents evidence that suggests that cortex may be involved in processing visual information in blindsight. A recent study has demonstrated that in visual cortex, frequency locking, especially between different visual areas, is difficult without feedback between areas that are connected. Removing striate cortex, with its massive reciprocal connections with several secondary cortical visual areas that are in turn further connected with each other and with additional visual areas, is likely to disrupt or even destroy this delicate phase locking.

2.1

INTRODUCTION

Although neurologists have often disagreed about the nature of visual disorders, they were for a century almost unanimous in agreeing that partial destruction or denervation of part of the striate cortex renders the patient clinically blind in the resulting visual field defect. The minority view, that some sensitivity remained or returned, was easily dismissed on the grounds that the visual cortex was merely damaged, not destroyed, or had partially recovered from damage. Despite the blindness, assessed by visual field perimetry, more recent investigations show that some patients with an apparently totally blind field defect, can detect and localize visual stimuli when persuaded to guess (Pöppel et al., 1973; Weiskrantz et al., 1974; Perenin & Jeannerod, 1978; Stoerig & Pöppel, 1986). Their high level of performance contrasts with their surprise when it is pointed out to them. Detection and discrimination of movement (Weiskrantz, 1986; Barbur et al., 1980; Magnusson & Mathiesen, 1989), flicker (Barbur et al., 1980; Weiskrantz, 1986; Blythe et al., 1987; Magnusson & Mathiesen, 1989) and orientation (Weiskrantz et al., 1974; Perenin, 1978; Weiskrantz, 1987) may also be present. Two patients asked to reach for solid objects, adjusted their grasp so that it matched the shape and size of the unseen objects. Even more surprisingly, they could use the meaning of unseen words flashed in their field defects in order to select between pairs of words subsequently presented in the intact visual field, i.e. they showed a priming effect (Marcel, 1983 a,b). These far-ranging residual visual capacities can be demonstrated even though the patients assert that they do not see anything in the field defects and are surprised when their often excellent performance is pointed out to them. This phenomenon has become known as ‘blindsight’ (Weiskrantz et al., 1974; Weiskrantz, 1986, 1990 for reviews) and has attracted the attention of philosophers, psychologists and neuroscientists because it highlights the nature of covert vision, indicates that striate cortex is indispensible for visual awareness, and provides a means of studying the visual information carried by pathways other than the major route through striate cortex.

2.2 THE VISUAL PATHWAYS

The pathways from the eye to their first projection zones in the brain are shown schematically in Fig. 2.1. Although most texts on vision describe the massive retinal projection to the dorsal lateral geniculate nucleus (dLGN) and to the superior colliculus (SC) the others are rarely mentioned, despite the fact that many of them, as judged from electrophysiological recordings, transmit information about the position, size, and movement of visual stimuli (e.g. Simpson, 1984). Given that several of them appear to deal with reflexive non-cognitive responses, e.g. the ventral lateral geniculate nucleus in the detection of light levels and intensity discrimination (Legg & Cowey, 1977a,b) the olivary pretectum in the control of the pupil, or the three accessory optic nuclei together with the nucleus of the optic tract in the detection of self-motion and the subsequent postural adjustments to flow fields (see Simpson, 1984 for review) they might seem excellent candidates for at least some aspects of blindsight. For instance, the change in skin conductance that follows the sudden presentation of a light persists when the light appears in the blind field (Zihl et al., 1980). The pupil continues to respond to changes in pattern and contrast, (Weiskrantz, 1990). And a patient with complete cortical blindness continued to follow a large moving striped display with his eyes, displaying optokinetic nystagmus, despite disclaiming any visual sensation that might have explained his visual tracking (Ter Braak et al.,