1st Joint meeting of

The Institute for Cognitive Science (ISC), Bron
&
The Institute of Cognitive Neuroscience (ICN), London

From perception to action representation :

Neural bases and disorders

25-26 septembre 2000

Organised by D. Boussaoud & P. Haggard
 

ISC, 67 Boulevard Pinel, 69675 Bron cedex
Phone: 33 (0)4 37 91 12 61; Fax: 33 (0)4 37 91 12 10
Contact: Driss Boussaoud (boussaoud@isc.cnrs.fr)

Monday
Tuesday
Monday abstracts
Tuesday abstracts
Discussion Groups
Posters abstracts

From perception to action representation :
Neural bases and disorders

Monday, september 25

12:00 Registration

13:45 Opening and welcome

Attention and space representation
Chair: D. Wolpert

14:00 P. Haggard (ICN, London) Awareness, binding and causation

14:30 J-R Duhamel (ISC, Bron) Neuronal basis of space representation

15:00 Y. Rossetti (INSERM U534, Bron) Several different “vision for action” systems

15:30 Coffee

Perception and action
Chair: P. Haggard

16:00 M-T Perenin (INSERM U371, Bron) Perception and action in unilateral neglect

16:30 A. Sirigu (ISC, Bron) The parietal cortex and the representation of action

17:00 L. Parsons (San Antonio, Texas) New Findings on the Role of the Cerebellum in Perception and Action.

17:30 C. Tallon-Baudry and O. Bertrand (INSERM U280, Lyon) Gamma oscillations and object representation in humans

18:00

Chair: T. Shallice
18:30 Plenary lecture by M. Jeannerod
Internal simulation of action as a unified framework for motor cognition

19:30 Session ends
20:00 Diner

From perception to action representation :
Neural bases and disorders

Tuesday september 26

Cognitive functions and their disorders
Chair: P. Jacob

9:00 T. Shallice (ICN, London) Fractionation of  the Executive systems

9:30 C. Frith (ICN, London) Consciousness and the functions of the prefrontal cortex

10:00 N. Georgieff (Hôpital Vinatier & ISC, Bron) Agency and its dysfunction

10:30 M. Saoud (Hôpital Vinatier, Bron) Context processing and action in schizophrenia

11:00 Coffee and poster viewing

11:30

12:00 Lunch

Modelling and psychophysics
Chair: C. Prablanc

14:00 J. Driver (ICN, London) Polysensory attention

14:30 Y. Paulignan (ISC, Bron) Perceptual awareness versus action

15:00 D. Wolpert (ICN, London) Computational human sensorimotor control

15:30 P. Burgess (ICN, London) The cognitive and neuroanatomical correlates of multitasking

16:00 Coffee

From perception to action representation : Neural bases and disorders

16:30 Discussion Groups
 Titles

ACTION THEME
Visuomotor adjustments: differences between motor performance and awareness of action
Helen Johnson and Patrick Haggard

The coding of movements in primary motor cortex: a TMS study
A.F. Hamilton, K.E. Jones, Z. Ghahramani*, R.N. Lemon & D.M. Wolpert

State estimation under stochastic and deterministic time-varying contexts
Philipp Vetter & Daniel M. Wolpert

Sensorimotor integration of retinal and gaze signals during smooth pursuit eye movements
Robert J. van Beers1, Daniel M. Wolpert2, Patrick Haggard1

Prism adaptation for different starting arm postures
Pierre Baraduc and Daniel Wolpert

Reaching and grasping in the monkey: effect of perturbation of object size and location.
A.C. Roy, Y.Paulignan, D.Boussaoud.

Neuronal activity related to bimanual coordination in the motor and premotor cortices of rhesus monkeys.
C. Jouffrais, E.M. Rouiller. Institute of Physiology. University of Fribourg

SPATIAL THEME
Is the Bell Inequality Violated in Visual Perception?
Andrew Duggins, Geraint Rees, Chris Frith

Vision and touch through the looking glass: cognitive remapping of multisensory information in the normal and damaged brain.
Angelo Maravita1, Charles Spence 2, Clair Sergeant 1, Karen Clarke 1, Masud Husain, & Jon Driver 1

Two systems of spatial representations in human posterior parietal cortex
S. Clavagnier*, I Schindler, Y. Paulignan, A. Vighetto, MT. Perenin

Convergent and divergent effects of neck proprioceptive and visual motion stimulation on ego- and allocentric space in neglect
Igor Schindler1, Georg Kerkhoff²

Spatial attention and memory vs motor preparation. The involvement of the premotor cortices as assessed by fMRI.
Stéphane Simon*, Martine Meunierº, Anna Berardi*, Loÿs Piettre*, Christoph Segebarth*, Driss Boussaoudº

Beware and be aware: Capture of attention by emotional stimuli in patients with hemispatial neglect
Patrik Vuilleumier (1,2) & Sophie Schwartz (1,3)

Impaired visual search and extinction following reversible inactivation of monkey lateral intraparietal area (LIP).
Claire Wardak and Jean-René Duhamel

Non-informative Vision Enhances Tactile Acuity
Steffan Kennett, Marisa Taylor-Clarke & Patrick Haggard

Predicting the future position of a looming target in 3D space: the role of retino-, head- and trunk-centered reference frames in man.
Marco Neppi-Mòdona*°, David Auclair,° Jean-René Duhamel°, Angela Sirigu°
* Facoltà di Psicologia, Università di Torino, Italia; °Institut des Sciences Cognitives, Bron, France

EXECUTIVE THEME
Agency judgement in schizophrenic patients.
C. Farrer , N. Franck , N.Georgieff , M. Marie-Cardine, J. Daléry, T. d ’Amato, & M. Jeannerod

In search of a monitoring deficit of action in schizophrenia : some arguments for a revision of this assumption.
Fourneret P 1., Franck N 1.2., Georgieff N 1.2., Jeannerod M 1

Can you remember the same way twice? Changes in the structure of autobiographical episodes recalled repeatedly at a short interval.
Laure Coates & Paul Burgess

Important deficit of action generation in schizophrenia: two experimental studies
Posada, N. Franck, N. Georgieff & M. Jeannerod.

Modelling the Somatoform and Dissociative (Conversion) Disorders
Richard J. Brown, PhD.

Predicting the sensory consequences of action
S-J. Blakemore, D.M. Wolpert & C.D. Frith

Planning and action knowledge in schizophrenic patients: Comparisons with patients with frontal lobe damage.
Tiziana Zalla, Andres Posada, Nicolas Franck, Nicolas Georgieff, Angela Sirigu
 

18:00 Meeting ends
 

From perception to action representation :
Neural bases and disorders
Monday Abstracts
Awareness, binding and causation

Patrick Haggard, Institute of Cognitive Neuroscience, UCL

The association between our intentions, our actions, and the effects these have on our environment are a key achievement of the human mind.  To achieve the status of a conscious agent, my mental processes must mark the external events caused by my intentional actions.  We call this process "efferent binding".

This talk will consider the subjective timing aspects of efferent binding.  Several converging lines of evidence have shown that the perceived time of discrete actions often precedes the onset of actual muscular movement.  We shall argue that psychological representations of actions arise partly from neural premotor processing.

We now report two new experiments on the perceived time of stimuli (beeps) and actions (keypresses) which occur in contingent pairs.  In the first experiment, we asked subjects to estimate the perceived time of stimuli and of actions in separate blocks.  The contingent relation between stimulus and action could run in either direction according to condition: either the stimuli elicited actions (in an SRT task), or the actions elicited by stimuli (in an operant task).  Subjects' estimates of stimulus timing and of action timing were compared with the perceived times of stimuli or actions made in isolation.  The method of Libet et al. (1983) was used to obtain timing judgements.  The biasing effects on the perceived time of stimuli and of actions of the additional event in the SRT and operant tasks were considered.

Briefly, we found strong perceptual attraction effects between percepts of the two events.  These are interpreted as evidence for efferent binding effects processes that influence conscious awareness.  In experiment 1, the causal event in each contingent task was perceived to occur at its normal time, whereas the consequent event was biased towards the cause.  Thus, in SRT, stimuli were perceptually fixed in time, whereas actions were perceptually attracted towards them.  In operance, actions were perceptually fixed in time, whereas stimuli were perceptually attracted towards them.  Experiment 2 compared timing judgements between the contingent tasks studied in the first experiment, and non-contingent, sequential tasks (two stimuli occurring in succession, or two actions made in succession).  Only contingent tasks showed the pattern of fixed causes and conflated effects.

These results suggest a specific perceptual attraction effect in the perceived time of events occurring in our basic interactions with our world.  It applies equally to sensory events and to motor events, and applies equally to operance and reaction.  Perceptual attraction may narrow the window of psychological time over which the mind must construct associations, and may allow effects to be individuated by their causes.  This could facilitate the process of efferent binding, and could thus underlie our sense of agency.

Acknowledgements: supported by MRC, BBSRC, DAAD and MPG.
 

Neuronal basis of space representation

Jean-René Duhamel,
Institut des Sciences Cognitives, CNRS, France

The parietal lobe contains multiple specialized areas which contribute to sensory analysis and movement preparation. Single cell recording evidence and the analysis of anatomical connectivity patterns in non-human primates reveal both similarities and differences in the core functional characteristics of each of the currently recognized parietal subdivisions. One of the main common features is that parietal areas integrate multiple sources of signals which are combined, presumably through population coding, in order to provide the spatial coordinates of directed actions.  Investigation of one parietal subdivision, the ventral intraparietal area (VIP),  demonstrates in particular that the activity of multisensory neurons is influenced by eye position in the orbit and that their receptive field, i.e. the region of the environment for which such neurons demonstrate spatial selectivity,  is encoded in coordinates which can be intermediate between a retinal and a head-centered reference frame. Neural network modeling suggests that such a complex form of coding could serve to perform multidirectional coordinate transformations across different sensory and motor modalities. Preliminary evidence from human psychophysical experiments will also be presented in support of the concept of multiple reference frames for encoding spatial locations.
 

Several different ‘vision for action’ systems

Yves Rossetti and Laure Pisella
Espace et Action, Institut National de la Santé Et de la Recherche Médicale, Unité 534
16 avenue Lépine, Case 13, 69676 Bron - France

There is a well-established argument for a double dissociation between vision for action and vision for conscious identification1. However, the complex neuroanatomical networks involved in vision, perception and action does not allow for a strict segregation between two cortical visual pathways, since numerous direct and indirect interconnections and convergence can be observed between the dorsal and the ventral stream2. Accordingly, evidence for a double interaction between the two visual systems have been documented. The residual sensori-motor performance of blindsight and numbsense patients is fully disrupted when a verbal representation of the stimulus is activated. This cognitive ? motor interference suggests that action can rely on the semantic representation (that is missing in these patients)3. Reciprocally, sensori-motor adaptation to a visual shift has been shown to affect cognitive representation of space in both neglect patients and normals4. This longer term sensori-motor ?cognitive interference suggests that higher-level representations of space can be re-organised by the plastic modification of sensori-motor correspondence.
The best way to isolate the sensori-motor mode of vision is to apply time constraints to the response being made by normal subjects. During a pointing experiment, subjects have been requested to immediately stop their action in case of a target jump. For speeded movements, they nevertheless performed numerous (counter-intentional) corrections toward the secondary target position. This automatic movement control was disrupted in a patient with a bilateral posterior parietal lesion . The posterior parietal cortex appears as an automatic pilot able to track a selected spatial target and drive the hand to it irrespective of the subject’s own intention5. The most typical function of the dorsal stream appears to be the on-line control of an ongoing goal-directed action.
It is concluded that, depending on the time scale considered, no interaction, or two way interactions between the dissociated vision for action and vision for identification can be observed6. The temporal dimension appears as one of the keys to the understanding of complex interconnected networks such as the visual brain.

1. Milner A.D., Goodale, M.A. 1995. The visual brain in action (Oxford Psychology Series 27). Oxford: Oxford University Press. 248pp.
2. Rossetti Y., Pisella L., Pélisson (2000) Eye blindness and hand sight: temporal aspects of visuo-motor processing. Visual Cognition, 7, in press.
3. Rossetti Y. (1998) Implicit short-lived motor representation of space in brain-damaged and healthy subjects. Consciousness and Cognition, 7, 520-558.
4. Rossetti Y., Rode G., Pisella L., et al. (1998) Prism adaptation to a rightward optical deviation rehabilitates left hemispatial neglect. Nature, 395, 166-169.
5. Pisella L, Gréa H, Tiliket C, et al. (2000) An automatic pilot for the hand in the human posterior parietal cortex toward a reinterpretation of optic ataxia. Nature Neuroscience, 3, 7, 729-736
6. Pisella L., Rossetti Y. (2000) Interaction between conscious identification and non-conscious sensori-motor processing: temporal constraints. in: Rossetti & Revonsuo (eds): Beyond dissociation: Interaction between dissociated implicit and explicit processing. Benjamins Amsterdam. pp. 129-151.
 

Perception and action in unilateral neglect

Marie-Thérèse Perenin
Cerveau et Vision, INSERM Unité 371, Bron
 

 The multiform aspect of the neglect syndrome has often been ascribed to the several ways space perception and representation may be altered. Some authors have stressed the prevalent role of a displacement and/or distortion of an egocentric frame of reference while other have pointed out a disturbance of object-centred coordinates or even of world-centred coordinates. Variations according to the prevalence of either a ‘perceptual’ or a ‘motor’ component have also been described in a number of studies. However, in most of them, non-natural, incompatible tasks were used, which prevent a clear interpretation of the results.
 In the present study, by recording natural, reaching and exploration hand movements, we have shown evidences for perception/action dissociations in both neglect patients and normal subjects with a neglect-like syndrome following optokinetic or vestibular stimulation. Despite a misperception of their sagittal midplane, all subjects performed virtually as normals when they had to point rapidly at visual targets. On the contrary, tactile exploratory movements and 'indirect’ pointing (after a delay or at a distance from the visual stimulus, i.e. in a different plane) were both influenced by the shift of the subjective straight ahead.
 These results provide further arguments for the hypothesis of two separate systems of spatial representations. ‘Motor’ representations dedicated to action are short-living, unconscious  and effector-specific processes; they require a strict temporal and spatial correspondence between the goal to be reached and the response directed at it. ‘Perceptual’ representations  are long-lasting and more elaborate processes which do not require such a stimulus-response correspondence. They can be memorized and give rise to delayed motor responses, or to responses at a distance from the stimulus, or to verbal responses concerning the perceived, explicit, localization of the stimulus. They would also take part in building a supramodal and conscious representation of space, on which rely deliberate exploratory behaviours.
 

The parietal cortex and the representation of action

Angela Sirigu

Institute for Cognitive Science, CNRS, Lyon, France.

 For over a century, the posterior parietal regions have been known to be involved in arm and hand movements,  from reaching and grasping movements to symbolic gestures and the manual use of tools and objects. Although the nature of this contribution is not fully understood, a recurring hypothesis is that the parietal cortex maintains some aspects of the internal models of the learned limb postures and motor programs  necessary for the guidance of skilled hand movements. In support of this hypothesis, I will present data: (1) on different patients with parietal cortex lesions who show selective impairments in mental movement rehearsal, using  a variety of tasks which can be used to establish the parallelism, or lack thereof, between imagined and executed movement sequence (Sirigu et al. , Neuroreport 1995, Sirigu et al. Science 1996; (2) recent fMRI results obtained in normal subjects who were asked to either imagine or to execute auditory-cued hand movements. Direct comparison between the two experimental conditions showed that specific cortico-subcortical areas were more engaged in mental simulation, including bilateral premotor, supplementary motor and left posterior parietal areas thus suggesting that specific brain areas are more involved and weigh more importantly on imagined movements rather than executed movements (Gerardin, Sirigu et al., in press, Cerebral Cortex, 2000).

Gamma oscillations and object representation in humans
C. Tallon-Baudry, O. Bertrand and C. Fischer.
INSERM U 280 Lyon

We experience objects as entities irrespective of whether they are perceived by our sensory system or recalled from memory. Since parts and properties of objects are encoded in many distinct brain areas, how does a coherent percept emerge? Rhythmic synchronization of neural discharges in the gamma band (around 40 Hz) could provide a spatial and temporal dynamic link between and within the areas involved in the same network, thus solving the so-called "binding problem" for object features [1]. We suggest that this mechanism could be used more generally for the construction of an object representation, driven either by sensory inputs or by internal top-down processes. If this hypothesis is correct, gamma-band oscillatory activity should appear in any task requiring the activation of an object representation.
Significant enhancements of induced activity (e.g. activity appearing with a jitter in latency from one trial to the next) in the gamma-band were repeatedly observed in human scalp EEG [2], in different tasks involving the activation of an object representation (a feature binding task, an hidden-object detection task, a delayed-matching-to-sample task). In addition, the different topographies of the induced gamma-band oscillatory activity suggest that depending on the task to be performed, different functional areas can be recruited to participate in an oscillatory ensemble. Intra-cranial EEG recordings in epileptic patients supported this interpretation: they revealed the existence in the extrastriate cortex of both within- and between-area oscillatory synchronization. Both local and long-distance synchronizations were task-dependent, and could account for the results obtained at the scalp level.
Scalp and intra-cranial recordings in humans thus suggest that the neural correlate of the activation of an object representation is the synchronization of a distributed cell assembly in the gamma-band.

1- Singer, W. & Gray, C.M. Annu. Rev. Neurosci. 18, 555-586 (1995).
2- Tallon-Baudry, C. & Bertrand, O. Trends Cogn. Sci. 3, 151-162 (1999)

From perception to action representation :
Neural bases and disorders

Tuesday Abstracts

Fractionating the Supervisory System

Tim Shallice

Institute of Cognitive Neuroscience, UCL.

A theoretical framework for fractionating the Supervisory System within the Norman/Shallice model of executive functions will be derived from the AI work of Fox and Dhas.  How adequately the framework relates to the subfunctions of regions of the frontal cortex will be considered from the prespectives of two types of empirical evidence. One type are functional imaging studies of the control processes involved in encoding and retrieval of episodic memories.  The second are neuropsychological studies of problem-solving and intention realisation.

Consciousness and the functions of prefrontal cortex

Chris Frith

Wellcome Department of Cognitive Neurology, Institute of Neurology, UCL.

Awareness of selecting one from a number of possible actions and then initiating that action forms an important component of consciousness. This awareness of selection arises especially for novel situations in which it is not immediately obvious which action is most appropriate. There is considerable evidence, from lesion and imaging studies, that dorso-lateral prefrontal cortex (DLPFC) has a major role in response selection in these circumstances. I propose that this selection is achieved by biasing an arbitrary, but appropriate subset of responses (sculpting the response space). If this is the case then the final initiation is achieved by some bottom-signal and not directly by activity in DLPFC.

It is not clear whether awareness is required for this kind of selection and initiation or whether activity in DLPFC is always associated with awareness. Electrophysiological studies suggest that awareness of selection is associated with the precise specification of the response to be made. Since this occurs rather late in the selection process controlled by the DLPFC, I would conclude that activity in DLPFC is not sufficient for awareness of response selection. Imaging studies directly examining awareness of selection implicate medial frontal structures including anterior cingulate cortex. Awareness of initiation precedes actual initiation and is therefore likely to be based on predicted rather than actual sensory consequences of action. The supplementary motor area (SMA) seems to have a critical role in the initiation of an action. This is probably achieved by inhibiting action until the appropriate moment. Thus SMA may have a critical role in the timing of awareness of action initation.
 

Context processing and action in schizophrenia

Mohammed Saoud
Hôpital Le Vinatier, Lyon

Patients with schizophrenia present a wide variety of deficits in the realm of cognitive functions. There is a challenge to understand such schizophrenia cognitive deficits in terms of a common unifying hypothesis. A single unifying hypothesis has difficulty explaining such disparate deficits, and, consequently, reference has been made to a variety of underlying models of normal cognitive functioning. One of the most coherent suggestions relates to impairments in context processing (extraction, representation and/or use of contextual information) (Servan-Shreiber et al., 1996). Such impairments were often explored in high levels of cognition, for instance, memory or language. Aside from the presence of various abnormalities, movement disorder is one characteristic of schizophrenia that is often described regardless of the theoretical framework used to define the disease. Context processing may apply to very simple pointing task as well as complex cognitive tasks. We will present several studies with different task difficulties ranging from simple speed-accuracy trade-off for target size to delayed pointing in a spatio-temporal context. Our results show that while the simplest context levels are processed normally in patients with schizophrenia, the extraction and use of more complex contextual information is impaired even in the case of goal directed action. We suggest that analysis of a low level of cognition, such as the natural tendency to adapt pointing movement to the task difficulty, could provide the basis for understanding higher level processing.

Perceptual awareness versus action ?

Y Paulignan, A.C. Roy and M. Jeannerod
Institut des Sciences Cognitives, CNRS UMR 5015, 67 boulevard Pinel,F69675 Bron cedex

In 1991 Castiello et al. presented an experiment showing a dissociation between a motor response to a perturbation and the corresponding awareness of the same event. Estimation of the delays was made by a double task paradigm : a motor and a vocal responses to the same stimuli. The results showed a large difference between latencies of motor and vocal (conscious) responses. The possible interactions between these tasks was a matter of debate.
The aim of the present study is to replicate this experiment with only one task.
Three objects were placed on a table at -10, 0 and 10 degrees of subject's sagittal axis. These objects were translucent dowels 10 cm height, 1.5 cm in diameter. In 80% of the trials the illumination of the central dowel was the signal for the subject to grasp it. On the remaining 20% of trials, the illumination was moved from the central object to one of the others at the beginning of hand movement. Movements were recorded by an OPTOTRAK system. After each perturbed trial the subjects were asked to show where the hand was when they detected the perturbation. This hand location was recorded. We computed the minimum of distance between the reported position and the wrist trajectory during the preceding perturbed trial. Then, we obtained a point on the wrist movement trajectory of the perturbed trial. This point determines the moment at which the subjects became aware of the perturbation.
Subject's awareness initiation occurs 410 ms after the perturbation. The motor response occurs later than for the previous studies of Castiello et al. (1991) and Paulignan et al. (1991).
The results confirm the Castiello's estimations of time of consciousness by giving the same values. They demonstrate that having to recall accurately the previous motor acts, hence being aware of them, has an influence on the motor performance.

Castiello, U., Paulignan, Y., & Jeannerod, M. (1991) Temporal dissociation of motor responses and subjective awareness : a study in normal subjects. Brain, 114, 2639-2655

Paulignan Y, MacKenzie CL, Marteniuk RG, and Jeannerod M (1991) Selective perturbation of visual input during prehension movements. 1. The effects of changing object position. Exp Brain Res 83: 502-512

Computational human sensorimotor control

Daniel Wolpert

Sobell Department of Neurophysiology, Institute of Neurology, UCL, UK.

The talk will focus on the computations underlying planning, prediction and learning in sensorimotor control.  First, I will present our work on motor planning which provides a unifying theory of eye and arm movement control. This theory suggest that trajectories are selected so as to minimize the consequences of signal-dependent noise. I will then present work on predictive internal models, neural circuits that predict the consequences of motor commands, focusing on their uses, how they are learned and their neural basis.  Finally, I will describe work on modular leaning in which we examine how humans learn to generate accurate and appropriate motor behaviour under many different and often uncertain environmental conditions.  This work focuses on a novel model, the multiple paired predictor-controller model, to deal with such selection and learning.
 

The cognitive and neuroanatomical correlates of multitasking

Paul W. Burgess
Institute of Cognitive Neuroscience, UCL, London

One of most curious groups of patients are those who are normal or even supra-normal on traditional neuropsychological tests, including those sensitive to frontal lobe damage, but who show deficits in planning, organisation and prospective memory in everyday life. These patients show a circumscribed deficit in situations that involve multitasking: i.e. situations that involve dovetailing different activities when returns to task are not directly signalled. This suggests that there are cognitive resources that are dedicated to this function.
This view receives support from a recent study of the Six Element Test (SET; the most commonly used clinical test of multitasking originally described by Shallice and Burgess, 1991). The carers of 92 mixed aetiology neurological patients were asked to rate the patients using a questionnaire measuring 20 of the commonest symptoms of the dysexecutive syndrome. factor analysis of these ratings revealed 5 orthogonal factors, with factor 2 related specifically to planning and organisation in everyday life. When correlations between the factor scores and the patients' performances on a range of clinical tests were examined, the scores for factor 2 showed only significant correlations with the SET, and not with WAIS-R IQ or a range of memory, language and other tests.
Further support is found in Burgess et al (2000). This study involved a componential analysis of a complex multitasking test using 60 patients with circumscribed cerebral lesions and 60 age- and IQ-matched controls. The tests was given in such a way that the relative contributions of rule learning, planning, plan following, autobiographical and retrospective memory to overall task performance could be assessed. Analysis of relations between lesion sites and test performance showed that damage to left medial posterior regions, including the posterior cingulate and occipital lobe, caused deficits on all aspects of the procedure except planning. Circumscribed impairments were however found in other lesion groups. Lesions to the right dorsolateral prefrontal cortex caused isolated planning impairment. The left anterior cingulate and surrounding regions were damaged in people who showed problems in remembering the task rules after a delay; and poor performance on the multitasking component of the test (measured as task switches minus rule-breaks) were associated with lesions to the left frontal pole.

Analysis of the relations between the different performance stages using structural equation modeling suggested that three primary are involved in multitasking. The first is retrospective memory, which is involved in rule learning and remembering. The second is required for planning, and the third is involved in the prospective memory demands of the task, such as remembering to switch tasks, and to follow one's plan.
Most recently, this methodology has been applied to the original version of the SET, with similar results, highlighting the role of anterior polar and superior medial regions in task switching after filled delays. Furthermore, two recent PET studies of this function have shown rCBF changes in this region during the maintenance of a delayed intention. The results could not be explained as increases in attentional demands. These results of this series of studies converge on the conclusion that there is a circumscribed set of cognitive processes that support human multitasking, and that the most likely supporting structures for the most dedicated processes are located in the medial and polar prefrontal regions.
 

From perception to action representation : Neural bases and disorders
Posters
Abstracts
 
Pierre Baraduc and Daniel Wolpert S-J. Blakemore, D.M. Wolpert & C.D. Frith Richard J. Brown
S. Clavagnier*, I Schindler, Y. Paulignan, A. Vighetto, MT. Perenin Laure Coates & Paul Burgess Andrew Duggins, Geraint Rees, Chris Frith
C. Farrer , N. Franck , N.Georgieff , M. Marie-Cardine, J. Daléry, T. d ’Amato, & M. Jeannerod Fourneret P., Franck N. , Georgieff N., Jeannerod M.  A.F. Hamilton, K.E. Jones, Z. Ghahramani, R.N. Lemon & D.M. Wolpert
Helen Johnson and Patrick Haggard C. Jouffrais, E.M. Rouiller. Steffan Kennett, Marisa Taylor-Clarke & Patrick Haggard
Angelo Maravita, Charles Spence, Clair Sergeant, Karen Clarke, Masud Husain, &  Jon Driver Marco Neppi-Mòdona, David Auclair, Jean-René Duhamel, Angela Sirigu A. Posada, N. Franck, N. Georgieff & M. Jeannerod
A.C. Roy, Y.Paulignan, D.Boussaoud Igor Schindler, Georg Kerkhoff Stéphane Simon, Martine Meunier, Anna Berardi, Loÿs Piettre, Christoph Segebarth, Driss Boussaoud
Robert J. van Beers, Daniel M. Wolpert, Patrick Haggard Philipp Vetter & Daniel M. Wolpert Patrik Vuilleumier, Sophie Schwartz
Claire Wardak and Jean-René Duhamel Tiziana Zalla, Andres Posada, Nicolas Franck, Nicolas Georgieff, Angela Sirigu

Prism adaptation for different starting arm postures
Pierre Baraduc and Daniel Wolpert

Sobell Dpt of Neurophysiology, Institute of Neurology, London

Prism adaptation is generally viewed as the learning of a congruence between two perceptual dimensions (Harris 1965; Bedford 1993). In accordance to this view, some authors found that the adaptation transfers from one type of movement to another  (Yachzel and Lackner 1977; Freedman et al. 1965). This view of adaptation as an alignment problem has been more recently challenged with the discovery that the magnitude of prism adaptation depends on the kinematics of  the pointing movement  (Kitazawa 1997).
These aftereffects had been formerly described as a sum of three components: (1) a visual shift; (2) a change in felt limb position; (3) an “assimilated corrective response”, that depends on the experimental context (Welch 1974). This third component has however not been properly characterised, and could correspond to the learning of a new sensorimotor coordination. It would therefore logically depend on all the parameters of the movement. This idea was checked by manipulating the initial posture of the arm.

Methods
Subjects were required to point with their right hand from a fixed starting point to a static target. The orientation of the arm at the beginning of the movement was restricted to a set of three postures that differed by the degree of adduction (see figure; for clarity, only 2 postures have been represented). During the pre-test period, movements with vision were interleaved with movements without vision. The visual feedback was only given when the starting position was the most adducted (pos. A). It was limited  to the representation of the finger endpoint. A horizontal visuomotor discrepancy was then progressively introduced, that reached a maximum of 10 cm. This discrepancy varied as a function of  arm extension, as a prism shift. During this exposure phase, the starting position was always A. Finally, a post-test phase identical to the pre-test assessed the amount of adaptation.

Results and discussion

These are preliminary results. Maximum adaptation is achieved for the starting position used during exposure. The magnitude of the aftereffects decreases sharply as a function of the initial abduction, from 80% adaptation for pos. A to 40% adaptation for pos. C. This suggests that a major component of the rapid visuomotor remapping is the acquisition of a new sensorimotor coordination. In accordance to this, a modelling study showed that two distinct learning mechanisms could be necessary to explain the whole range of aftereffects (Guigon, personal communication): (1) a corrective supervised learning, analogous to the correction based on visual feedback alone; (2) a slower Hebbian mechanism, analogous to the realignment between proprioception and vision.
As the end posture varies as a function of the starting posture, it cannot however be excluded that these results are due to a difference in final proprioception. The variability of final arm posture does not lend a strong support to this hypothesis. Nevertheless, a control experiment will have to also constrain the final orientation of the arm.

References
Bedford F (1993) in The psychology of learning and motivation pp 1-60.
Freedman SJ, Hall SB, Rekosh JH (1965) Percept Motor Skills 20:1054-1056.
Harris CS (1965) Psychol Rev 72:419-444.
Yachzel B and Lackner J (1977) Percept Psychophys 22:147-151.
Welch RB (1974)  J Exp Psychol 103:700-705.
 

Predicting the sensory consequences of action

S-J. Blakemore, D.M. Wolpert & C.D. Frith

Wellcome Department of Cognitive Neurology, University College London

We have investigated how the brain distinguishes between self-produced and externally produced sensations. 'Forward models' predict the sensory feedback from self-produced movements thereby enabling us to recognise the sensory consequences of our own actions. It is proposed that an impairment of this predictive mechanism might give rise to certain symptoms experienced in schizophrenia (1,2). If self-produced sensations are interpreted as being generated by an external source, then thoughts might be interpreted as external voices (auditory hallucinations) and self-produced movements might be interpreted as externally generated (delusions of control or passivity phenomena). Psychophysical studies have shown that self-produced tactile stimulation is rated as less intense and 'tickly' than externally produced stimulation, and we have proposed that such an attenuation of self-produced tactile stimulation is due to the sensory predictions made by a forward model. Functional neuroimaging studies have demonstrated that such attenuation is mediated by somatosensory cortex and the anterior cingulate cortex: these areas are activated less by a self-produced tactile stimulus than by the same stimulus when it is externally produced. Furthermore, evidence suggests that the cerebellum might be involved in generating the prediction of the sensory consequences of movement (3). Finally, we have recently demonstrated that psychotic patients with auditory hallucinations and passivity phenomena show no attenuation of self-produced sensory stimulation (4). This supports the proposal that these symptoms are associated with an impairment of the functioning of the forward model.

1. Frith, CD. (1992). The Cognitive Neuropsychology of Schizophrenia. Lawrence Erlbaum Associates, UK.
2. Frith, CD, Blakemore S-J & Wolpert, DM. Abnormalities of the Perception and Control of Action. In Press in Philosophical Transaction of the Royal Society of London: Biological Sciences
3. Blakemore, S-J, Wolpert, DM & Frith, CD (1998). Central cancellation of self-produced tickle sensation. Nature Neuroscience  1(7), 635 - 640
4. Blakemore, S-J, Smith, J, Steel, R, Johnstone, E & Frith, CD. The perception of self-produced sensory stimuli in patients with auditory hallucinations and passivity experiences: Evidence for a breakdown in self-monitoring. In press in Psychological Medicine.

Modelling the Somatoform and Dissociative (Conversion) Disorders
Richard J. Brown, PhD.
Institute of Neurology, UCL.

Traditional models have emphasized the role of motivational factors in the pathogenesis of the Somatoform and Dissociative (conversion) disorders. However, such factors are absent in many instances of somatoform and dissociative illness. Based on an integrative model of cognition, somatoform and dissociative symptoms are reconceived here as distortions in subjective experience arising from the chronic activation of symptom representations during the generation of consciousness. By this view, somatoform and dissociative illness are essentially cognitive phenomena rather than the product of emotional trauma or the expression of unconscious conflict as is traditionally conceived. Several factors that moderate the occurrence of these conditions via their effect on the activation of symptom representations are identified, including symptom monitoring, the activation of complementary representations, high-level rumination and low-level attentional sensitivity.

Two systems of spatial representations in human posterior parietal cortex
S. Clavagnier*, I Schindler, Y. Paulignan, A. Vighetto, MT. Perenin
INSERM U 371, ISC, 69500 Bron, Hôpital Neurologique 69003 Lyon, France

Although often associated, spatial disorders from parietal origin may occur in isolation. This is the case for optic ataxia and unilateral neglect, which result from lesions respectively centered on the dorsal and the ventral part of posterior parietal cortex. This neurological dissociation as well as psychophysical findings has led to the hypothesis of two separate systems of spatial representations. "Motor" short-lived are dedicated to action and subtend goal-directed movements. "Perceptual", long-lasting representations are dedicated to space perception; they can be memorized and used in motor responses at a distance from the stimulus or on exploratory behavior. These two kinds of representations would be subserved by the superior and inferior parietal lobules respectively.
Optic ataxia patients would thus be improved when required to perform either delayed or displaced responses, while neglect patients would perform worse. In the present study, visuo-motor performances of two patients, one with optic ataxia (IG) and another one with unilateral neglect (FP), were compared to that of normal control subjects. Subjects were required to point at visual targets presented randomly in their peripheral field. Pointing responses of the right hand were recorded by using an optoelectronical motion analyzer (OPTOTRAK?) in three conditions: "direct" pointing, "indirect" pointing with either a delay of five seconds between stimulus offset and movement onset, or a spatial separation between stimulus and responses planes. While normal subjects and FP pointing movements became less accurate in the “indirect” pointing conditions, disturbances of IG performances decreased significantly in these conditions. These results indicate that optic ataxia patients still have access to a cognitive mode of spatial representations, the one that is impaired in neglect patients. This new dissociation confirms the idea of a functional subdivision in posterior parietal cortex.
 

Can you remember the same way twice? Changes in the structure of autobiographical episodes recalled repeatedly at a short interval.

Laure Coates & Paul Burgess
Institute of Cognitive Neuroscience, UCL (University College London)

Using verbal protocol analysis of healthy participants' recollections, Burgess and Shallice (1996) proposed a model of the processes involved in remembering events that have happened to you. They found that in addition to memories themselves, participants produced many protocol elements that were evidence for the operation of cognitive control processes in autobiographical recollection. The model proposes three broad categories of control process. Descriptor processes are assumed to produce a specification of the type of trace that would satisfy the demands of the retrieval task. Memory Editor processes are involved in checking that the output fits with previously retrieved memory elements in the episode being retrieved and also with the overall task requirement. Mediator processes control strategic and problem-solving operations concerning the adequacy or plausibility of retrieved memory elements.

If the operation of these control processes corresponds to the process of recollection rather than the products of it, then subsequent recollection of them should be weaker than for the memories that were recalled. We therefore investigated the proportional change in control elements when people are asked to recall the same autobiographical episode twice. 16 participants were asked 12 questions about everyday events that happened to them, such as "Describe the last time you had dealings with the police". Immediately after they finished answering all 12 questions, they were asked half of these questions again, and had to repeat exactly what they said the first time they answered them.

The results showed a significant proportional decrease in all control elements (Editor, Mediator and Descriptor) in the second recollection episode, with a corresponding proportional increase in the recollection of memories. This pattern even remained when subjects were asked not merely to recall the episode again, but also when they were asked to recall the exact words they had used in giving their first answer. This pattern also appeared in recollection of other people’s verbal protocols. The results support the hypothesis that cognitive control processing in recollection has a different status from the activation of memory representation.
 

Is the Bell Inequality Violated in Visual Perception?

Andrew Duggins, Geraint Rees, Chris Frith

Wellcome Department of Cognitive Neurology, University College London

The perceptual latency of a direction change in an array of oscillating squares was shown by Moutoussis and Zeki to be significantly greater than the perceptual latency of a colour change . This has led to the suggestion that there is a ‘microconsciousness’ generated at each of several anatomically separate and functionally specialised nodes . Furthermore, anatomical evidence suggests that there is relative segregation of inputs to the different nodes. From these theories follows a further experimental prediction, that the probability of perceptual pairing of colour and motion oscillations must not violate the Bell Inequality : We will review a standard derivation of the Inequality, and explain how it can be applied to the Moutoussis/Zeki paradigm. Surpisingly, the results of the original experiment suggest that the Inequality is violated, and thereby cast doubt on the ‘microconsciousness’ theory.  Alternatively, it is possible that the critical assumption of perceptual pairing, which would allow application of the Bell Inequality in this context, is not valid. In a new experiment, we replicate the Moutoussis/Zeki paradigm in a way that allows these two alternative interpretations to be resolved, and which allows statistical test of the ‘microconsciousness’ theory. Even isolating those trials in which perceptual pairing is confirmed by subject report, we find that the Inequality is violated. We conclude that the activity of cells in V5 must indeed depend partly on colour input and the activity of cells in V4 partly on motion. Visual consciousness must then be considered non-local and inseparable: the microconsciousness does not exist.

Agency judgement in schizophrenic patients.

C. Farrer , N. Franck , N.Georgieff , M. Marie-Cardine, J. Daléry, T. d ’Amato, & M. Jeannerod .

One of the central questions about the problem of self-consciousness and other-consciousness is to understand the experience of agency or how a subject attributes an action to himself or to an other agent. One approach for understanding the underlying mechanisms is to study disturbances of self-consciousness in schizophrenic patients. Indeed, one class of symptoms displayed by these patients, the so-called positive symptoms (e.g: delusion of control, hallucinations, ...)  are suggestive of an alteration of the awareness of one’s own action and of the recognition of actions performed by others.
One hypothesis postulates that an attribution of action relies on a discrimination between central representations activated from within and those activated by external cues (Georgieff and Jeannerod, 1998). A first experiment  by Daprati et al (1997) has tempted to study attribution of action in a group of schizophrenic patients. The results have shown that schizophrenic patients with hallucinations and delusion of control tended to overattribute to themselves actions produced by others and in conditions where the cues for discriminating the origin of an action were degraded.
In order to analyse more precisely the influence of the perceptive information on this agency judgement  we realised another experiment where the parameters of the visual information  were controlled using an electronic device. Subjects hold a joystick with their right hand. The movements of the joystick  were fed into a virtual hand holding a joystick. The virtual image was projected on a mirror overlying the subject’s hand. This device allowed us modifying the apparent direction and/or velocity of the movement actually performed by the subject. A manual motor task and an agency judgement about this task were performed by 29 normal subjects and 29 schizophrenic subjects
The results obtained in schizophrenic patients showed  a clear deficit for all the patients when their movements were delayed up to 250-280ms. Only the influenced patients were severely impaired for angular biases up to 30°. Those results could not be explained by deficits such as perceptual or attentional mechanisms impairment, indeed all the patients  performed well at the BORB which tests those abilities. An impairment in the  detection of visual cues indicating the intention of the movement (as it has been shown in our group of influenced schizophrenic patients) can lead to  a misunderstanding of  the action of an agent. So a deficit at this level could explain the fact that influenced patients tended to self-attribute movements they observed. This result is reinterpreted in the theoretical framework of Georgieff and Jeannerod.
 

Daprati E, Franck N, Georgieff N, Proust J, Pacherie E, Dalery J, Jeannerod M, (1997) : Looking for the agent: an investigation into consciousness of action and self-consciousness in schizophrenic patients. Cognition; 65 : 71-86.
Georgieff N. and Jeannerod M, (1998). Beyond consciousness of external reality: a “who” system for consciousness of action and self-consciousness. Consciousness and Cognition; 7: 465-477
 

In search of a monitoring deficit of action in schizophrenia : some arguments for a revision of this assumption

Fourneret P 1., Franck N 1.2., Georgieff N 1.2., Jeannerod M 1
1 : Institut des Sciences Cognitives. 67 Bd Pinel – 69675 Bron
2 : CHS Le Vinatier. 95 Bd Pinel – 69677 Bron

Abstract

The first-rank symptoms (Schneider, 1959), among which delusions of control, thought insertion, verbal hallucinations, are certainly the most impressive clinical signs met in schizophrenia and the most detrimentalto the social functioning of the patients. According to Frith (2000), these symptoms result from a lack of awareness of certain aspects of motor control ; this deficit is, itself, in relation with a failure in the cognitive mechanism by which the predicted consequences of an action can be derived from a forward model based on the intended sequence of motor commands. This assumption of an impairement in the central monitoring of their own actions by the schizophrenic patients, is derived from several experimental studies, which highlighted a greater difficulty for the patients, compared with controls, of being able to correct erroneous movements in the absence of visual feedback (Malenka, 1982 ; Frith & Done,1989 ; Mlakar et al, 1994).
In our study, 19 schizophrenic patients (10 with Schneiderian symptoms and 9 without Schneiderian symptoms) versus 19 paired control subjects  were subjected to a sensorimotor adjustment task. Using a magnetic stylet, they had to connect two visual targets with a trajectory as straight as possible. The subjects could not see their hand during the test and could not see the result of their action before the last third of the distance separating the
two targets. In the experimental condition, the computer generated a conflict between the action planned and the sensory-motor feedback by a linear directional bias (15°) to the right. Thus, to succeed the task, subjects had to modify their motor program and deviate their hand by 15° in the opposite direction. All subjects were naive at the beginning of the test and performed twenty trials. Sensory-motor adjustment to the bias was evaluated by the surface between the line traced and the ideal line to compensate for the deviation (square medium ratio). Awareness of the conflict was appreciated by the explicit report of a left manual correction.
Our results show that only the patients aware of the manual correction (n= 9) succeeded the task with a performance not significantly different from the control subjects. Nevertheless and surprisingly, most of them presented Schneiderian symptoms. For the others (mainly patients without Schneiderian signs), although the degree of adjustment was significantly lower than in controls, it showed evidence for a broadly preserved central monitoring of action. Only the ability to become aware of the motor intention appeared to be defective in these subjects.
These results do not support the assumption of a deficit in central monitoring of action in schizophrenic patients, especially those suffering from Schneiderian symptoms. Furthermore, they reinforces the idea, defended by some authors, of a functional dissociation between the consciousness of action and the executive control of action.

The coding of movements in primary motor cortex: a TMS study

A.F. Hamilton, K.E. Jones, Z. Ghahramani*, R.N. Lemon & D.M. Wolpert

Sobell Dept of Neurophysiology, Institute of Neurology
*Gatsby Computational Neuroscience Unit.

Whether movements are represented in Cartesian, joint, muscle or in some other co-ordinate system in primary motor cortex is still unclear. Almost all evidence comes from studies carried out in non-human primates. To examine movement coding non-invasively in humans we have used Transcranial Magnetic Stimulation (TMS) over the arm area of motor cortex to produces a force pulse generated by the muscles in the arm.  While the coil remained stimulating the same area of M1, the arm was moved over a range of joint angles covering the workspace. We recorded the force vectors at the wrist and the EMG in four muscles of the upper arm produced by the TMS pulse. The array of force vectors and EMG over the workspace showed a systematic pattern of change. We will present preliminary results and discuss their implications for the coding debate.
 

Visuomotor adjustments: differences between motor performance and awareness of action

Helen Johnson and Patrick Haggard

Institute of Cognitive Neuroscience, UCL.

Previous studies using the double-step pointing paradigm have suggested that visuomotor adjustments to a target shift can be dissociated from conscious perceptual awareness of the target shift.  This study investigated the action awareness subjects have of their own visuomotor adjustments, by asking subjects to repeat in their own time the movement paths made in immediately preceding double-step pointing trials.

Subjects were asked to reach out and touch a central illuminated LED as quickly and as accurately as possible.  On 33% of trials the LED ‘jumped’ 10 cm to left or right 25 ms after movement onset.  Subjects were instructed before each block to respond to the target jump in one of four ways, according to condition:
1) Point: if the LED jumped the subject had to follow it
2) Antipoint: if the LED jumped the subject had to point to the equivalent opposite location
3) Overpoint: if the LED jumped, the subject had to point to a location twice the distance of the jump in the same direction
4) Overantipoint: if the LED jumped, the subject had to point to a location twice the distance in the opposite direction
In all conditions subjects simply pointed to the central LED if it did not jump. Immediately after completing each trial subjects repeated the path of their original movement as accurately as possible.  The LED remained at its final location throughout this repetition period.

Visuomotor adjustments were more rapid in the basic point condition than in the other conditions.  We then compared the original movements with the repetitions to measure the extent to which subjects were aware of the path of their original movement, and in particular, their awareness of any visuomotor adjustments.  In the basic point condition subjects believed that they began to adjust later and to a lesser extent than they actually did.  In the antipoint condition, in contrast, they overestimated the extent of their adjustment. That is, they thought they deviated further in the intended direction, opposite to that of the LED, than they actually did.  In the overpoint condition subjects initially underestimated the extent of their adjustment, as in the basic point condition, and then, later in the movement, overestimated it, as in the antipoint condition. In the overantipoint condition subjects overestimated only the end of the deviation.

These findings provide evidence that visuomotor adjustments may be dissociated from awareness of action.  When the visuomotor adjustment directly corresponds to the perceptual target shift, as in the basic point condition, visuomotor adjustment is mediated by an automated response system of which subjects are at least partly unaware.  Thus, subjects believe that they performed basic double-step adjustments slower and less accurately than they actually do.  In the other conditions the required adjustment requires some indirect transformation of the perceptual target shift.  These adjustments are mediated by a slower, controlled response system that not only enters awareness but is also slightly exaggerated, leaving subjects believing that they performed the task better than they actually did.  Interestingly, the overpoint condition may reflect an amalgamation of the two systems, in which an automated initial response to follow the LED is subsequently modified.

Acknowledgement: Helen Johnson is supported by a BBSRC Special Committee Studentship.  Additional funding was provided by the Royal Society.
 

NEURONAL ACTIVITY RELATED TO BIMANUAL COORDINATION IN THE MOTOR AND PREMOTOR CORTICES OF RHESUS MONKEYS

C. Jouffrais, E.M. Rouiller.
Institute of Physiology. University of Fribourg

As movements of the two arms are naturally coupled, there must be a neural mechanism which can possibly influence this bimanual coupling. To investigate the role of motor cortical areas in the control of bimanual spatial and temporal coordination, we recorded single unit activity in the primary motor and premotor cortices of a monkey trained to execute a conditional reach and grasp drawer task. Four different delayed movements -unimanual right, bimanual right-left (reaching with the right hand, grasping with the left hand), unimanual left or bimanual left-right (reaching with the left hand, grasping with the right hand) - were instructed depending on the color of a visual cue. The gaze position of the monkey was controlled using the scleral search coil technique. This design allowed us to examine the signal-, set- and movement-related activity of individual cells involved in specific unimanual or bimanual movements. We classified the cells in three different categories possibly contributing to the coding of bimanual movements: (1) the “combination neurons” exhibited a spike rate modulation during bimanual movements which was the combination of the discharges observed during unimanual movements; (2) the “invariant neurons” showed signal- and/or set-related activity which was the same whatever the task the monkey was performing; (3) the “interaction neurons” exhibited discharge rate modulations which specifically reflected the interaction of the two arms during at least one of bimanual tasks. These preliminary results suggest a possible role for premotor and motor cortices in bimanual coordination.
 

Non-informative Vision Enhances Tactile Acuity
Steffan Kennett, Marisa Taylor-Clarke & Patrick Haggard
Institute of Cognitive Neuroscience, UCL (University College London), UK.

Previous work has revealed interdependence between the visual and tactile systems. For example, vision and touch appear to be intimately linked during both reflexive and voluntary shifts of spatial attention [1, 2]. Furthermore, single-unit recording has shown multimodal cells with spatially congruent receptive fields in both modalities [3]. Sight of a hand has been shown to reduce detection times to tactile targets presented to it [4]. However, no previous work has investigated whether vision affects the informational content of tactile signals, and in particular the spatial resolution of the tactile map of the body surface. Such evidence would add weight to suggestions that crossmodal links may arise due to feedback from populations of multimodal neurons to primary sensory cortex [e.g., 5]. Such feedback may modulate receptive field size or even lead to short-term cortical plasticity. Previous behavioural studies cannot distinguish between crossmodal modulations in the salience of a tactile stimulus (which could arise at any of several stages in processing pathway) and modulations in informational content (acuity), which are likely to arise at the level of the primary cortical projection. In this study, we directly measured effects of vision on tactile acuity by using a staircase procedure to estimate tactile two-point discrimination thresholds (2PDTs) on the arm. Participants always gazed towards their arm throughout the tactile stimulation session but, via use of a half-silvered mirror, the visual scene they viewed was varied across four conditions. In the first condition, participants gazed at their arm in total darkness (DARK condition). In the second condition, participants had clear vision of the stimulated part of their arm, except during a 700ms interval when the tactile stimulators were moving to strike the arm (VISION OF ARM condition). In the third condition, participants saw a cylindrical object at the location of the stimulated part of the arm (VISION OF OBJECT). In the fourth condition, participants viewed their arm through a 2.5? magnifying glass (MAGNIFIED VISION OF ARM). The dark interval around the time of tactile stimulation in conditions 2 and 4 ensured that vision could not be directly informative for tactile discrimination judgements.

Tactile acuity was improved in the VISION OF ARM condition relative to both DARK and VISION OF OBJECT conditions. The latter difference rules out spatial attention to the stimulated location as the sole explanation of the improved performance. Interestingly, acuity was further improved in the MAGNIFIED VISION OF ARM condition relative to the VISION OF ARM condition. This provides the first direct evidence that crossmodal links modulate somatosensory spatial representations.

Figure:  Schematic view of the experimental set-up in VISION OF ARM condition. Participants placed their right arm inside a box that had a semi-silvered mirror in one wall (pale shading). When lights inside the box were illuminated the arm was clearly visible, but the retracted tactile stimulators (tactors) were hidden by the opaque walls (dark shading). Lights were extinguished during tactile stimulus delivery (either one or two stimuli) to preclude any informative vision. (The tactors are shown at the point of contact with the arm; they would not have been visible at this position during the experiment.) Participants wore a patch over their right eye.

1. Spence, C., Nicholls, M.E.R., Gillespie, N. & Driver, J. (1998). Perception & Psychophysics, 60, 544-557.
2. Spence, C., Pavani, F. & Driver, J. (in press). Journal of Experimental Psychology: Human Perception and Performance.
3. Graziano, M.S.A. & Gross, C.G. (1993). Experimental Brain Research, 97, 96-109.
4. Tipper, S.P., Lloyd, D., Shorland, B., Dancer, C., Howard, L.A. &  McGlone, F. (1998). Neuroreport, 9, 1741-1744.
5. Hahnloser, R., Douglas, R.J., Mahowald, M. & Hepp, K. (1999). Nature Neuroscience, 2, 746-752.
 

Vision and touch through the looking glass: cognitive remapping of multisensory information in the normal and damaged brain.

Angelo Maravita1, Charles Spence 2, Clair Sergeant 1, Karen Clarke 1, Masud Husain, &  Jon Driver 1

1 -Institute of Cognitive Neuroscience, University College London
2 -Department of Experimental Psychology, University of Oxford
3 - Imperial College School of Medicine, Charing Cross Hospital, London

Integration of visual and tactile stimuli is particularly effective when visual stimuli are delivered near to tactually stimulated body parts. Studies on monkeys show that response of cross-modal visual-tactile neurons in several regions of the cortex is stronger for visual stimuli presented in the vicinity of the hands. This makes functional sense, as related visual and tactile information will normally come from similar locations, near the skin surface. However, there is one common situation in which stimuli on our body produce the visual appearance of being placed far away. This happens when we observe ourselves in mirrors, while acting on our own body (shaving, making-up, etc.). Tactile-proprioceptive cues then signal stimuli on the body, while visual information suggests that related visual stimuli are placed at a distance (“through the looking glass”).
We have shown in both normal subjects, and in a right-hemisphere patient with crossmodal extinction, that  visual information delivered close to the hand, but observed only via its mirror reflection in far space, is treated by the brain as a stimulus near the body in peripersonal space.

Normal subjects were tested in a visual-tactile interference paradigm. Crossmodal interference with tactile judgements is usually stronger for visual distractors close to the tactually stimulated hands, than for visual distractors in far space. In one of our conditions (Mirror) visual distractors were delivered from lights near the hands, but the lights and hands could only be observed as reflections in a mirror placed in front of the subjects. In another condition (Box), the half-silvered mirror was illuminated so as to act like a window into the  box on which it was mounted. Subjects now observed the contents of the box, where visual distractors and two stuffed rubber hands were placed far away from the subject, at the exact positions consistent with the reflections of visual distractors and subject’s hands in the Mirror condition. Interference from visual distractors on tactile judgements was stronger in the Mirror than the Box condition. We therefore suggest that visual afference from the mirror, although optically appearing as far away (“through the looking glass”) is computed by the normal brain as being near to the corresponding body part. A similar finding was observed in a neurological case for whom right visual events could extinguish awareness of tactile events on the left hand, especially if the visual event was close to the right hand. Such crossmodal extinction was stronger for a distant right visual stimulus in the Mirror condition, than in the Box condition.
These results suggest that mirror reflections projecting the image of distant visual events, can be recoded as arising from peripersonal rather than extrapersonal space.

Predicting the future position of a looming target in 3D space: the role of retino-, head- and trunk-centered reference frames in man.

Marco Neppi-Mòdona*°, David Auclair,° Jean-René Duhamel°, Angela Sirigu°
* Facoltà di Psicologia, Università di Torino, Italia; °Institut des Sciences Cognitives, Bron, France.
 

Avoiding or intercepting looming objects implies not only a precise estimate of time to contact but also of impact location. We present an experiment designed to investigate the relative contribution of retino-, head-, and trunk-centered reference frames in a trajectory prediction task. Normal subjects have to predict, in complete darkness, the impact location on their face (left or right of their head midline) of a LED fixed to a robotic arm moving in 3D space with a looming trajectory originating from a leftward, straight-ahead or rightward spatial location. Manipulating subjects' gaze and trunk orientation, produced important and systematic errors in the prediction of impact locations, resulting in a strongly biased estimation of subjective impact in a direction ipsilateral to the hemispatial origin of the trajectory. We demonstrate that predicting the future position of a moving target 1) implies access to a multiple reference frames system and 2) the spatial alignment of the reference frames among them and with respect to the origin of the trajectory is critical for a correct prediction. Furthermore, normal subjects' performance was not homogeneous, some individuals relying mostly on retinal cues, others on head-centered cues and yet others on mixed spatial cues. These results are consistent with current concepts of space representation in the parietal cortex, which stress the simultaneous coding of visual information in multiple reference frames.
 

Action generation in schizophrenia: two  experimental studies
A. Posada, N. Franck, N. Georgieff & M. Jeannerod.
Institute for Cognitive Science - CNRS, 67 Bd Pinel, 69675 Bron, France.

We can define a ‘mind level’ where words, or symbols, acquire mean, are consciously managed, can be communicated to others through a language and can generated voluntary actions. We call this level ‘explicit‘ by its strong relation with language. Our intentions, which will guide our actions to their completion are always generated at this level and one of their characteristics is they are ahead of execution. We used this characteristic to design the first experiment and evaluate the ability of schizophrenics to generate precise actions based on  explicit information. In this task subjects explicitly learned a repetitive color sequence and then received the instruction to give an anticipatory motor response by pushing corresponding color keys before the next element in the sequence was displayed on a computer screen. Different types of sequences (temporal and spatial) and experimental conditions were tested in both a group of normal subjects and a group of schizophrenic patients. Schizophrenia is a condition known to alter conscious executive function. Our results showed a strong deficit in performing the anticipation task in schizophrenic patients. Although they were found to be able to acquire a conscious knowledge of the sequences almost normally, their anticipation ability was reduced by comparison to normal subjects in all the tested conditions. After this positive result, we realized a second experiment to evaluate more precisely the deficit of schizophrenics to perform these particular actions. In this task, we measured reaction time in three different conditions: simple reaction to a color displayed on screen, categorization of the color displayed on screen (push corresponding color button) and implementation of an explicit rule to modify the color-button relation of the second condition. As control subjects did, schizophrenic patients showed similar pattern response in the first and the second condition; the reaction times of patients were a few slower. But in the third condition when subjects used the explicit rule, the reaction times of patients grew considerably. And, unlike the controls which showed a progressive diminution of reaction times through time (reflecting an automation of rule), schizophrenics showed an enhancement of the reactions times (reflecting fatigue and the absence of any automation). These results are interpreted as a deficit of schizophrenic patients in consciously using the explicit information to consequently  generate actions and create motor programs. They also expand the notion of a working memory deficit in schizophrenia and bear strong implications for understanding executive disorders observed in such patients.
 

Reaching and grasping in the monkey: effect of perturbation of object size and location.
A.C. ROY, Y.PAULIGNAN, D.BOUSSAOUD.
Institut des Sciences Cognitives, BRON, FRANCE

Human prehension movements are classically described as consisting of two components: a reaching component guided by spatial location of objects, and a grasping one guided by their size and shape. To understand the anatomical and physiological correlates of these visuomotor channels, we sought to describe the kinematics of reaching and grasping in macaque monkey trained to reach for and grasp 3-D objects. In this study, we tested the degree of independence of reaching and grasping by perturbing either object location or its size. Hand path and grip aperture were measured in three dimensions using an optotrak system.
Two animals are included in this study, and the analysis was based on 8 000 movements. Both monkeys were tested on perturbation of object location and size. Perturbation of object location affected both reaching and grasping components. In particular, grip aperture displayed a double peak pattern in 20% to 80% of trials, depending on the direction of perturbation. By contrast, perturbation of object size lead to a single peak of grip aperture, corresponding to the last object size. This observation indicates that the new object size has been taken into account earlier. As it has been reported in human subjects, size perturbation induces a smooth reorganization of movement  and "recoordination" of reaching and grasping. These findings suggest that reaching and grasping are not strictly independent visuo-motor channels. This view contrasts with observations of segregated anatomical pathways linking parietal cortex and premotor areas.
 

Convergent and divergent effects of neck proprioceptive and visual motion stimulation on ego- and allocentric space in neglect

Igor Schindler1, Georg Kerkhoff²

1INSERM U 371"Cerveau et Vision", 18 avenue Doyen Lépine, F-69675 Bron, France
²EKN - Clinical Neuropsychology Research Group, Dept. of Neuropsychology, Bogenhausen Hospital, Dachauerstr. 164, D-80992 München, Germany

Spatial neglect can affect egocentric as well as allocentric-based reference frames. The egocentric aspect of the disorder is manifested as an exploration bias corresponding to the ipsilesional deviation of the subjective visual straight ahead estimation (SSA). The allocentric deficits can be observed in the size estimation task as well as distance-judgment independently of the relative position of the stimulus configuration to the individuals head and body sagittal plane, thus attesting these patients a size- and space-distortion beyond their ipsilesionally deviated SSA. Whereas optokinetic or slow visual motion stimulation (SVS) as well as neck-muscle vibration have been shown to modulate essentially the egocentric aspect, effects on allocentric spatial deficits were only reported for optokinetic stimulation.
To this purpose we compared the modulatory effects of SVS and neck vibration on two types of visual-spatial tasks in neglect: One egocentric, measuring the visual subjective straight ahead (SSA) and three allocentric spatial tasks in the horizontal dimension: Line bisection, size and distance judgment. Five patients with left-sided spatial neglect were tested across four conditions: A baseline condition without any manipulation was compared with the experimental condition of SVS to the left and left neck vibration. As control condition we used vibration of the left hand and righward SVS.
In the baseline condition all patients not only showed a right-sided deviation of the visual straight ahead but also a left-sided size distortion in line bisection and size estimation as well as a space distortion for left-sided distance estimation. SVS to the left improved significantly the SSA as well as the size and distance estimation. In contrast during neck muscle vibration only the SSA was significantly ameliorated. In the control conditions no influence on the allocentric tasks was obvserved.
The present data suggests that SVS not only affects the representation of the subjective midline but also allocentric space representations. In contrast the efficacy of neck muscle vibration seems to be specific for egocentric tasks. One explanation is, that leftward background motion facilitates the direction of attention towards neglected regions of space. Another, compatible hypothesis, attributes the motion effect to a facilitating generation of egocentric space representation by providing a directional visual input to this disturbed representation. We propose that the largely intact visual motion system is capable of modulating the neuronal activity in the lesioned parieto-temporal cortex of neglect patients. These findings offer therapeutic applications for neglect rehabilitation since motion stimulation obviously exerts a powerful influence on perception.
 

The Role of Medial Prefrontal Cortex in the Representation of Task Specific Meaning

Sophie K. Scott, Alex Leff, Richard R. J. Wise.
Institute of Cognitive Neuroscience UCL.

Two positron emission tomography (PET) studies are presented which address the neural correlates of representations in different central executive tasks. A role is identified for left medial prefrontal cortex in the generation, representation and choice between alternative response mappings for individual stimuli. This system is involved in central processing of sound structure, or read words, when the task requires that the stimuli be assessed in a novel and ambiguous manner, in which multiple candidate responses are considered. Activity in this region correlates with longer reaction times in such choice tasks, but not in other tasks where the response class is not ambiguous.

Spatial attention and memory vs motor preparation. The involvement of the premotor cortices as assessed by fMRI.
Stéphane Simon*, Martine Meunierº, Anna Berardi*, Loÿs Piettre*, Christoph Segebarth*, Driss Boussaoudº
* INSERM U438, Grenoble, France. º Institut des Sciences Cognitives, Lyon, France

Introduction. Single cell recordings in awake behaving monkeys have shown that, in visually guided movements, the majority of neurons in the premotor cortex are preferentially active in relation to motor preparation, rather than to spatial attention or memory1. To examine whether these findings apply to the human brain, we used fMRI (ten healthy volunteers) combined with experimental paradigms very much inspired by those applied in neurophysiological studies in monkeys.
Material and methods. Paradigms. Block paradigms (8 min overall) were applied, comprising three control and three activation epochs of equal duration. Two paradigms were designed to dissociate activation related to motor preparation (MP) from that reflecting spatial attention and/or working memory (SAM). Subjects were asked to press one of two keys depending on the color of visual cues presented on a screen. Visual stimuli and motor responses were identical in the two conditions, but subjects performed different tasks. In the SAM paradigm, control and activation epochs differed in terms of attentional/mnemonic requirements, whereas in the MP paradigm they were different in terms of motor preparation. An example of trials performed during the control and activation epochs of the SAM paradigm is shown below (Fig 1). The central cross represents the fixation cross. White squares were presented successively at pseudo-random locations within the visual field (presentation time 500 ms, delay between successive presentations of white squares was pseudo-randomized, ranging between 1.2 and 2.0 s). Following presentation of a variable number of white squares, two adjacent red and green squares were presented for 1.5 s, and the fixation cross turned either red or green. One of these colored squares was at the same location as the last white square. During the activation epochs, subjects were instructed to attend to and memorize the successive positions of the white squares, and press one of the two keys depending on the color of the square positioned at the location of the white square. During the control epochs, the subject’s response was determined by the color of the fixation cross. Thus, during the control epochs, the performance of the task did not require spatial attention and/or memory.
MR acquisition and data processing. Examinations were performed at 1.5 T (Philips NT). A GRE EPI sequence was applied (TR=3.7s, TE=45ms, ?=90°). The volume of interest was composed of 25 adjacent slices. Resolution was (4 mm)3. Functional maps were generated by means of SPM 96. Statistical significance threshold for clusters obtained in the group analysis was established at p=.05.

Figure 1. SAM paradigm.

Results. SAM paradigm. A large activation was found, bilaterally, within the posterior superior parietal cortices. The right dorsolateral prefrontal cortex and, bilaterally, the lateral and mesial premotor cortices were also activated (Fig 2A). A faint activation was also found within the caudate nuclei.

Figure 2. Activations in premotor areas for the SAM (A) and MP (B) paradigms. Talairach y coordinates extend from -19 (top) to +9 (botttom). Right hemisphere is on right side.

MP paradigm. No activation was detected within the parietal cortices. Frontal activations were found, bilaterally, within the latero-dorsal and mesial premotor cortices, with extension into the left precentral gyrus (Fig 2B). Subcortically, activations were also obtained within the caudate nuclei, the right thalamus, the left substantia nigra and the red nucleus, as well as within the left amygdala.
Discussion and conclusion. An interesting finding is the different involvement of the premotor cortices for spatial attention/memory vs motor preparation, as illustrated in Figure 2. The posterior part of the mesial as well as lateral left premotor cortices are devoted to motor preparation. In contrast, the anterior part of the lateral premotor cortex seems specifically involved in spatial attention and memory. These observations are in line with recent studies and support the notion of functional distinction between anterior and posterior regions of the premotor cortex2-4.
 References
1. Boussaoud D et al., Behav Brain Res (1996), 72: 1-15.
2. Tanné J et al., NeuroReport (1995), 7: 267-272.
3. Johnson P et al, Cereb Cortex (1996), 6: 102-119.
4. Toni I et al, Cereb Cortex (1999), 9: 35-49.
 
 

Sensorimotor integration of retinal and gaze signals during smooth pursuit eye movements

Robert J. van Beers1, Daniel M. Wolpert2, Patrick Haggard1
1Institute of Cognitive Neuroscience, UCL (University College London), UK.
2Sobell Department of Neurophysiology, Institute of Neurology, UCL, UK.

To localise a seen object, the central nervous system has to integrate the object's retinal location with the direction of gaze. We investigated this process by examining the localisation of static objects during smooth pursuit eye movements. We performed a series of experiments in which subjects performed localisation tasks. From the localisation errors found in these experiments we could derive the errors in the retinal and in the gaze signals and also how these signals are combined.
During an eye movement, a static object's image moves across the retina. Objects that produce retinal slip are known to be mislocalised [1, 2]: objects moving toward the fovea are seen too far on in their trajectory, whereas errors are much smaller for objects moving away from the fovea. These effects are usually studied in experiments in which subjects localise the moving object relative to a briefly flashed one during fixation. In this situation the moving objects are mislocalised, but flashes are not. In our first experiment we found that a similar differential mislocalisation occurs for static objects relative to flashes in the retinally equivalent case during pursuit, in accordance with earlier results [1, 3]. In addition, we found that this effect is not specific for horizontal pursuit, but it was also found in other directions. In a second experiment we examined how this effect generalises to positions outside the line of eye movement. We found that large localisation errors were found in the entire hemifield ahead of the pursuit target, and these were predominantly aligned with the direction of eye movement. In a third experiment, we determined whether it is the flash or the static object which is mislocalised ahead of the pursuit target. Subjects tracked a target and compared the positions of two sequentially presented stimuli, the first one ahead of the pursuit target and the second one behind it. We found that when the first stimulus was a flash, it was mislocalised, but when it was a static object, it was not. The nature of the second stimulus did not have an effect. These results demonstrate that during pursuit it is the flash, not the static object, which is mislocalised. Note that this is the reversed pattern of that during fixation. In a fourth experiment, subjects pointed at flashed and static stimuli which were presented in complete darkness during pursuit. The results were similar to those of the previous experiment. This shows that the mislocalisation of flashes during smooth pursuit reflects absolute rather than relative localisation, and is therefore based on integrated retinal and gaze signals.
We conclude that the central nervous system compensates for the retinal localisation errors (for objects which produce retinal slip) during smooth pursuit eye movements in order to maintain position constancy for static objects. This compensation is achieved in the process of sensorimotor integration of retinal and gaze signals: different retinal areas are integrated with different gaze signals to guarantee the stability of the visual world.

1. Mateeff, S. and Hohnsbein, J. (1988) Vision Research 28: 711-719.
2. Mateeff, S., Yakimoff, N., Hohnsbein, J., Ehrenstein, W. H., Bohdanecky, Z., and Radil, T. (1991) Vision Research 31: 131-138.
3. Mitrani, L. and Dimitrov, G. (1982) Vision Research 22: 1047-1051.

State estimation under stochastic and deterministic time-varying contexts

Philipp Vetter & Daniel M. Wolpert

Sobell Department of Neurophysiology, Institute of Neurology, University College London

The consequences of a motor command depend on the prevailing movement context, that is the state of our own body and the environment. This context is not static but changes over time. For instance the weight of a bottle changes as we pour from it, and muscles fatigue with use. We explored how the central nervous system's estimate of finger position evolves when visual feedback is removed by introducing a discrepancy between the true and displayed finger position. In the first experiment the context changed stochastically between two discrete discrepancies, and in the second experiment the discrepancy changed continuously in a deterministic fashion.

In the stochastic experiment, subjects made sequential pointing movements, during which their visual feedback was offset from the finger by two possible fixed amounts. When visual feedback was removed, subjects' pointing tended toward that appropriate for the average of the two contexts. This behavior is optimal in the sense that it minimizes the mean squared pointing error.

In the deterministic experiment, the displayed finger height changed with respect to the true height in a sinusoidal fashion over time. Subjects were unaware of this perturbation, and continued to compensate for the time-varying sinusoidal discrepancy, even in the absence of visual feedback. Their performance demonstrates that subjects can estimate the rate of change of the context. Taken together, these results suggest that the central nervous system actively models both stochastic and deterministic components of the time-varying context without requiring awareness.

Beware and be aware: Capture of attention by emotional stimuli in patients with hemispatial neglect
Patrik Vuilleumier (1,2) & Sophie Schwartz (1,3)

(1) Institute of Cognitive Neuroscience, University College, London, UK
(2) Department of Neurology, University of California, Davis, CA, USA
(3) Institute of Cognitive Sciences, University of California, Berkeley, CA, USA

Adaptive-evolutionary arguments and empirical evidence suggest that emotional significance of stimuli might prioritize the allocation of spatial attention to potentially relevant stimuli. We asked whether such effects might still occur in patients with unilateral neglect and visual extinction who usually remain unaware of contralesional stimuli when these compete with concurrent ipsilesional stimuli. In a first experiment, shapes or faces with either neutral, happy, or angry expressions were presented in right, left, or both visual fields. On bilateral trials, three parietal patients extinguished faces on the contralesional side much less often than shapes, and faces with happy or angry facial expressions much less than neutral faces. In a second experiment, pictures of spiders or flowers made of similar low-level features were presented in right, left, or both fields. Again, parietal patients were more likely to perceive emotional stimuli (spiders) than other similar but neutral pictures (flowers). We suggest that in patients with neglect in whom mechanisms of spatial attention are impaired after parietal damage, intact visual pathways to the ventral temporal lobe and amygdala could still mediate mechanisms of "emotional attention".
 

Impaired visual search and extinction following reversible inactivation of monkey lateral intraparietal area (LIP)
Claire Wardak 1 , Etienne Olivier 2  and Jean-René Duhamel 1
1 Institut des Sciences Cognitives, CNRS, Bron, France
2 Université catholique de Louvain, Bruxelles, Belgique

Behavioral expressions of unilateral sensory and/or motor neglect have been studied experimentally in monkeys following lesions  restricted to the parietal cortex, to the frontal cortex and to certain subcortical structures. Severe and lasting deficits have rarely been observed in monkeys, as compared those described in humans following right hemispheric damage. This discrepancy has been interpreted as reflecting the absence of functional hemispheric lateralization in the monkey, which permits greater compensation of the deficits through neuronal plasticity mechanisms. Reversible inactivation methods overcome this problem by allowing the study of reproducible, acute behavioral effects in the same animal. We inactivated the lateral intraparietal area (LIP), a region of the parietal cortex implicated in visual attention and saccadic eye movements, by placing discrete injections of muscimol, a GABA agonist, at sites identified through single unit recordings, in a macaque performing visual search and saccades to simple and double simultaneous stimuli. No change in the latency or accuracy were observed during visually-guided saccades to single contralesional targets. However, the frequency of saccades to briefly flashed contralesional targets is reduced if a simultaneous ipsilesional target is presented. Also, the contralesional target was often undetected when presented alone. In the visual search task, feature search of a colored target among distractors of a different color is unaffected by LIP inactivation, but conjunction search of a target defined by both color and shape shows a marked contralesional impairment and a concurrent ipsilateral facilitation. These deficits show on saccade latency and on the proportion of trials where the initial saccade went to a distractor. Inactivation of the adjacent ventral intraparietal area (VIP) had no effects on any of these tasks.  These results indicate that LIP inactivation produces a selective deficit in attentional target selection, but no saccade programming or execution impairment. Selective reversible inactivation of the parietal cortex can thus produce behavioral impairments analogous to those observed in human neglect, and could be used to gain a better understanding of how the functional subdivisions of the parietal lobe contribute to the different aspects of this syndrome.

Planning and action knowledge in schizophrenic patients: Comparisons with patients with frontal lobe damage.

Tiziana Zalla, Andres Posada, Nicolas Franck, Nicolas Georgieff, Angela Sirigu

Institut des Sciences Cognitives, BRON, France
 

Schizophrenia is characterized by disorders in a large variety of cognitive domains including executive function, attention, memory, learning and language. A line of researches has drawn attention to a putative frontal dysfunction or a frontal dementia and documented that schizophrenics are impaired on executive deficits. However, the issue whether executive disturbances in schizophrenic patients correspond to selective deficits qualitatively comparable to the performance of frontal lobe patients is still controversial, mostly because standard neuropsychological assessments, such as the Wisconsin Card Sorting Test, do not allow to disentangle the component processes required for executing the task.
By using a set of multicomponent tasks which segregated various information processes used in action planning, we assessed the ability to generated  knowledge and elaborate a plan of action in a group of seventeen schizophrenic patients. Their performance was compared with that of a group of nine patients with frontal lobe lesions and a group of sixteen normal control subjects. Schizophrenics were found to perform significantly worse than normal subjects and frontal lobe patients in retrieving action knowledge data base related to non-routine activities. Moreover, similarly to frontal lobe patients, they were impaired in processing the sequential organization of actions in both the generation and the ordering tasks, in setting priorities among the script actions, in script rule violation, and in estimating the importance of the individual actions in relation to the script's goals. These results indicated that schizophrenics are impaired in processing critical features of action knowledge necessary to accomplish a plan of action and that this pattern of deficits is similar to that observed in patients with damage to frontal cortex.


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