Effect of distorted visual feedback on the sense of agency.

It has been hypothesized that an internal model is involved in controlling and recognizing one's own actions (action attribution). This results from a comparison process between the predicted sensory feedback of the action and its real sensory consequences. The aim of the present study is to distinguish the respective importance of two action parameters (time and direction) on such an attribution judgment. We used a device that allows introduction of discordance between the movements actually performed and the sensory feedback displayed on a computer screen. Participants were asked to judge whether they were viewing 1) their own movements, 2) their own movements modified (spatially or temporally displaced), or 3) those of another agent (i.e, the experimenter). In fact, in all conditions they were only shown their own movements either unaltered or modified by varying amounts in space or time. Movements were only attributed to another agent when there was a high spatial discordance between participants' hand movements and sensory feedback. This study is the first to show that the direction of movements is a cardinal feature in action attribution, whereas temporal properties of movements play a less important role.

[Frontal lobe, executive functions and cognitive control]. / Lobe frontal, fonctions exécutives et controle cognitif.

The approach of executive functions began with the early description of behavioural disorders induced by frontal damage. The development of neuropsychology has led to the description of a large variety of cognitive disorders. The interpretation in cognitive terms of these disorders has emphasized the critical impairment of executive functions which are involved in non-routine situations. The role of working memory, initially suggested by animal studies, has also largely influenced theoretical approaches. Numerous studies have been interpreted within the theoretical frameworks developed by Shallice and by Baddeley. A large amount of studies have allowed the specification of control processes (initiation, inhibition...). Recent studies are beginning to investigate cognitive disorders underlying behavioural changes such as deficits of emotional, social and metacognitive processes. In addition these studies seriously question the assumption of a unique central control system. These studies have deeply influenced the clinical approach, the assessment and the diagnosis of executive syndrome. For clinical practice, these data lead to favour specific assessment of some key behavioural and cognitive deficits based on a battery of tests and structured interview of an informant.

Grasping objects: the cortical mechanisms of visuomotor transformation.

Grasping requires coding of the object's intrinsic properties (size and shape), and the transformation of these properties into a pattern of distal (finger and wrist) movements. Computational models address this behavior through the interaction of perceptual and motor schemas. In monkeys, the transformation of an object's intrinsic properties into specific grips takes place in a circuit that is formed by the inferior parietal lobule and the inferior premotor area (area F5). Neurons in both these areas code size, shape and orientation of objects, and specific types of grip that are necessary to grasp them. Grasping movements are coded more globally in the inferior parietal lobule, whereas they are more segmented in area F5. In humans, neuropsychological studies of patients with lesions to the parietal lobule confirm that primitive shape characteristics of an object for grasping are analyzed in the parietal lobe, and also demonstrate that this 'pragmatic' analysis of objects is separated from the 'semantic' analysis performed in the temporal lobe.

Mental motor imagery: a window into the representational stages of action.

The physiological basis of mental states can be effectively studied by combining cognitive psychology with human neuroscience. Recent research has employed mental motor imagery in normal and brain-damaged subjects to decipher the content and the structure of covert processes preceding the execution of action. The mapping of brain activity during motor imagery discloses a pattern of activation similar to that of an executed action.

Mental imaging of motor activity in humans.

Motor imagery corresponds to a subliminal activation of the motor system, a system that appears to be involved not only in producing movements, but also in imagining actions, recognising tools and learning by observation, as well as in understanding the behaviour of other people. Recent advances in the field include the use of techniques for mapping brain activity and probing cortical excitability, as well as observation of brain lesioned patients during imaging tasks; these advances provide new insights into the covert aspects of motor activity.

Visual cognition: a new look at the two-visual systems model.

In this paper, we argue that no valid comparison between visual representations can arise unless provision is made for three critical properties: their direction of fit, their direction of causation and the level of their conceptual content. The conceptual content in turn is a function of the level of processing. Representations arising from earlier stages of processing of visual input have very little or no conceptual content. Higher order representations get their conceptual content from the connections between visual cognition and other parts of the human cognitive system. The two other critical properties of visual representations are their mind/world direction of fit and their mind/world direction of causation. The output of the semantic processing of visual input has a full mind-to-world direction of fit and a full world-to-mind direction of causation: it visually registers the way the world is and is caused by what it represents. The output of the pragmatic processing yields information for the benefit of intentions, which clearly have a world-to-mind direction of fit and a mind-to-world direction of causation. An intention is both the representation of a goal and a cause of the transformation of a goal into a fact. These properties segregate representations specialized for perception from those specialized for action. Perception implies comparison between simultaneously represented and analyzed objects: hence, object perception presupposes the representation of spatial relationships among objects in a coordinate system independent from the perceiver. Spatial relationships carry cues for attributing meaning to an object, so that their processing is actually part of semantic processing of visual information. These considerations lead to a re-evaluation of the role of the two classical pathways of the human visual system: the ventral and the dorsal cortical pathways. The parietal lobe, which has been identified with the dorsal pathway, cannot be considered as a unitary entity with a single function. The superior parietal lobule carries visuomotor processing, a non-lateralized process. The right inferior parietal lobule contributes to the perception of spatial relationships, a process with a mind-to-world direction of fit and a world-to-mind direction of causation. Finally, the left inferior parietal lobule contributes to still another type of representation, related to visually goal-directed action, i.e., with both a world-to-mind direction of fit and a mind-to-world direction of causation.

Defective recognition of one's own actions in patients with schizophrenia.

OBJECTIVE: The possibility that delusions of influence could be related to abnormal recognition of one's own actions was investigated in persons with schizophrenia. METHOD: Schizophrenic patients with (N=6) and without (N=18) delusions of influence were compared with normal subjects (N=29) on an action recognition task. The image of a virtual right hand holding a joystick was presented to the subjects through a mirror so that the image was superimposed on their real hand holding a real joystick. Subjects executed discrete movements in different directions. Angular biases and temporal delays were randomly introduced in some trials, such that the movement of the virtual hand departed from the movement executed by the subjects. After each trial, subjects were asked whether the movement they saw was their own. RESULTS: Compared with normal subjects, both patient groups made significantly more recognition errors in trials with temporal delays. In trials with angular biases, the error rate of patients with delusions of influence significantly differed from that of comparison subjects and from that of patients without delusions of influence. CONCLUSIONS: The findings support the hypothesis that delusions of influence are associated with a quantifiable difficulty in correct self-attribution of actions. This difficulty may be related to a specific impairment of a neural action attribution system.

Mechanisms of visuomotor coordination: a study in normal and brain-damaged subjects.

This paper reviews the role of vision in controlling pointing and reaching movements in man. Studies of visuomotor behaviour in normal subjects allow the identification of several levels for describing organization of visuomotor control. The relative contribution of central (programming) and peripheral (feedback) mechanisms; the role of the eye-head coordinated position as a reference for controlling accuracy and direction of movements; and the relative independence of input-output channels in controlling proximal and distal segments of the musculature. These levels represent a useful framework for understanding pathological disruption of visuomotor control produced by cortical lesions.

Mental imagery in the motor context.

The working hypothesis of the paper is that motor images are endowed with the same properties as those of the (corresponding) motor representations, and therefore have the same functional relationship to the imagined or represented movement and the same causal role in the generation of this movement. The fact that the timing of simulated movements follows the same constraints as that of actually executed movements is consistent with this hypothesis. Accordingly, many neural mechanisms are activated during motor imagery, as revealed by a sharp increase in tendinous reflexes in the limb imagined to move, and by vegetative changes which correlate with the level of mental effort. At the cortical level, a specific pattern of activation, that closely resembles that of action execution, is observed in areas devoted to motor control. This activation might be the substrate for the effects of mental training. A hierarchical model of the organization of action is proposed: this model implies a short-term memory storage of a 'copy' of the various representational steps. These memories are erased when an action corresponding to the represented goal takes place. By contrast, if the action is incompletely or not executed, the whole system remains activated, and the content of the representation is rehearsed. This mechanism would be the substrate for conscious access to this content during motor imagery and mental training.

Limited conscious monitoring of motor performance in normal subjects.

Normal subjects traced sagittal lines on a graphic tablet using a stylus held in their right hand. The hand was hidden by a mirror in which they saw the lines projected from a computer screen. In normal trials, the line seen in the mirror exactly corresponded to the traced line. In perturbed trials, a bias was introduced by the computer, so that the line appeared to deviate in one direction (right or left) by a variable angle (2, 5, 7 or 10 degrees). Subjects consistently displaced their hand in the opposite direction for producing a visually sagittal line. After each trial, they were asked in which direction they thought their hand had moved. In perturbed trials, they grossly underestimated the hand deviation. In addition, a post-hoc analysis revealed that one group of subjects misperceived the direction of their hand movement in the direction opposite to the perturbation (Group 1, including 9 Ss), whereas the other group gave responses in the correct direction (Group 2, including 4 Ss). In a second session using the same experimental paradigm, a motor response was asked for: subjects had to indicate the perceived direction of their hand during each trial by drawing a line with their eyes closed. Again, responses indicated a poor conscious monitoring of motor performance. These results suggest that normal subjects are not aware of signals generated by their own movements.

Impairment of grasping movements following a bilateral posterior parietal lesion.

The observation of a patient (A.T.) with a bilateral posterior parietal lesion of vascular origin is reported. A.T. presented a bilateral (more marked on the right) deficit in grasping simple objects (neutral cylindrical dowels) without deficit in reaching toward the location of these objects. The major symptom was an exaggerated anticipatory opening of the fingers with poor correlation with object size, resulting in awkward grasps. It was present both when the hand was visible to the subject and when it was not. This deficit was much less marked if the neutral objects were replaced by usual objects of the same sizes. Finally, in tasks where she had to indicate with her fingers the size of visual objects presented as virtual images through a mirror, or the size of imagined usual objects, A.T. performed normally. These results are discussed within the framework of a dual representation of objects. Only the "pragmatic" representation for steering object-oriented actions would be impaired in this patient as a result of posterior parietal damage. By contrast the semantic representation for object identification would be intact.

In search for the egocentric reference. A neurophysiological hypothesis.

Unilateral lesions of the middle suprasylvian gyrus in the cat parietal cortex produce an asymmetrical vestibulo-ocular reflex such that responses to head rotation toward the side opposite to the lesion become weaker. Unilateral lesions of the superior colliculus in the cat also produce the same effect. In either case symmetrical responses are recovered within 2-3 weeks. These results are discussed in terms of displacement, by the unilateral lesion, of an internal reference used for directing behaviour within extrapersonal space. Relevance of this hypothesis to clinical symptoms observed in man after unilateral posterior parietal lesions is discussed, with particular emphasis on the unilateral neglect phenomenon.

Analogical transfer is effective in a serial reaction time task in Parkinson's disease: evidence for a dissociable form of sequence learning.

Several studies of procedural learning in Parkinson's disease (PD) have demonstrated that these patients are impaired with respect to age-matched control subjects. In order to examine more closely the specific impairment, we considered three dimensions along which a procedural learning task could vary. These are: (1) implicit vs explicit learning, (2) instance vs rule learning, and (3) learning with internal vs external error correction. We consider two hypotheses that could explain the impairments observed in PD for different types of explicit motor learning: (H1) an impairment related to the acquisition of rules vs specific instances, and (H2) an impairment in learning when no explicit error feedback is provided. In order to examine the condition of rule learning with external error feedback, we developed a modified version of the serial reaction time (SRT) protocol that tests analogical transfer in sequence learning (ATSL). Reaction times are measured for responses to visual stimuli that appear in several different repeating sequences. While these isomorphic sequences are different, they share a common rule. Verbatim learning of a sequence would result in negative transfer from one sequence to a different one, while rule learning would result in positive transfer. Parkinson's patients and age-matched controls demonstrate significant acquisition and positive transfer of the rule between sequences. Our results demonstrate that PD patients are capable of learning and transferring rule or schema-based representations in an explicit learning format, and that this form of learning may be functionally distinct from learning mechanisms that rely on representations of the verbatim or statistical structure of sequences.

Is the organisation of goal-directed action modality specific? A common temporal structure.

With the help of kinematic analysis, the temporal organization of the complex daily activity 'drinking from a bottle with a glass' was described in detail. The analysis focused on the sequential action structure, the prehensile acts, and the bimanual coordination as well as on the effect of different instruction modalities on these parameters to explore the underlying representation for this complex action. Movements of the two arms were recorded in three-dimensional space with the help of an optoelectronic device in 12 normal subjects under four conditions: (1) action pantomime after verbal instruction; (2) action imitation after observation of the action performed by the experimenter without the objects; (3) action pantomime while seeing, but not touching the objects; and finally (4) action execution with objects. Despite high execution variability, the temporal structure of the action could be precisely described by the relative duration and peak velocity of action segments, by the MGA-object size-correlation, and by linear regression analysis between the onsets of functionally related action segments. A similar structure of the action as characterized by these kinematic parameters was retained across different instruction modalities. Only when the action was executed with the objects, the interval between the movement onsets of either hand and the peak velocity of the manipulative acts were reduced, while no change was observed across the other three instruction modalities. This stability of the temporal structure suggests the existence of a level in the representation of an action where all the modalities converge.

Motor imagery of a lateralized sequential task is asymmetrically slowed in hemi-Parkinson's patients.

We examined seven right-handed, asymmetrical (right side affected) Parkinson's disease patients and seven age-matched controls in a manual finger sequencing test using left and right hands in vision, no vision, and motor imagery conditions. All patients displayed motor asymmetry, favoring the left hand. They also displayed motor imagery asymmetry, mentally simulating movement more slowly with their right affected hand than with their left hand. Additionally, impairment in mental hand rotation correlated significantly with the imagery asymmetry. These data support two related hypotheses: (a) Motor sequence imagery and execution share common neural structures. (b) The frontostriatal system is among these shared structures.

The prefrontal cortex and conscious monitoring of action: an experimental study.

To investigate the role of the prefrontal cortex in conscious monitoring, we used an experimental paradigm generating a conflict between the action planned and the sensory-motor feedback. We analyzed the acquisition of explicit knowledge of the strategy for resolving the conflict and its influence on motor adaptation. Twenty patients with frontal lobe lesions and 18 controls had to trace a sagittal line with a stylus on a graphics tablet. A mirror on which the traced line, processed by a computer, was projected hid the hand. A mask limited visual feedback to the last third of the trajectory. Without informing the subjects, the line traced was modified by introducing a bias of 24 degrees to the right. To succeed in the task, subjects had to modify their motor program and to deviate their trajectory in the opposite direction. Conscious elaboration of the strategy was evaluated by the number of trials needed to explicitly report the required deviation. Three groups of patients were distinguished: (1). with normal explicit strategy; (2). with delayed explicit strategy, and (3). without explicit strategy at the last trial. They significantly differed by the severity of the dysexecutive syndrome, particularly of environmental adherence. Motor adaptation was evaluated by the area between the line traced and the ideal line to compensate for the deviation. In patients with normal elaboration of the strategy, motor control was similar to that of controls, but it was severely disturbed in the other two groups. These results suggest the involvement of the prefrontal cortex in conscious motor monitoring.

Visuo-motor control of the ipsilateral hand: evidence from right brain-damaged patients.

We investigated the extent to which the right hemisphere is involved in the control of the ipsilateral hand by analysing the kinematics of right-hand prehension in right brain-damaged (RBD) patients. We required patients to grasp one of five possible objects, equally-sized and distributed over a 40 degrees wide workspace. With the purpose of investigating the right hemisphere contribution to the on-line visuo-motor control, we also assessed patients' ability to correct their movement "in-flight", in response to a sudden change of object position. Patients' performance was compared to that of aged-matched controls. A Younger group of healthy subjects, matching the population classically tested on double-step paradigms, was also evaluated to fully assess whether patients' kinematics could be partially due to normal ageing. As a further aim, the possible influence of hemispatial neglect was evaluated by comparing the performances of right brain-damaged patients with and without neglect. In normal subjects, the results confirmed and extended the notion of (a). positional tuning of grip formation, and (b). fast reactions following a change in object position. In addition, subtle effects of ageing on visuo-motor behaviour were shown by less efficient movement correction in the Elderly group. Patients executing reach-to-grasp actions into the left contralesional hemispace were selectively affected in both temporal and spatial aspects of movements. While their performances were relatively well preserved in the right hemispace, patients did not show positional tuning of grip formation, nor fast corrections of their movements when acting in the left hemispace. Interestingly, similar deficits were found irrespective of the presence of neglect. These results show that the right hemisphere contributes to the processing of visuo-motor information that is necessary for executing actions with the ipsilateral hand in the contralateral space.

Effects of unilateral flocculus lesions on vestibulo-ocular responses in the cat.

Unilateral lesions of the cerebellar flocculus were performed in three chronically-implanted adult cats. Following the lesion a spontaneous nystagmus was observed in the dark, with the fast phase directed to the lesioned side. Vestibulo-ocular responses in the dark became asymmetrical. Responses to velocity steps exciting the labyrinth ipsilateral to the lesion were strongly increased. A decrease, although less marked, was observed in the opposite direction. Responses to sinusoidal oscillations in the dark were also asymmetric with respect to both the cumulative eye displacement during rotation in the two directions and the interval between two consecutive reversals of eye movement. These differences were greater at the lower tested frequencies (0.01 HZ) than at the higher (0.1 HZ). Spontaneous nystagmus disappeared in about 10 days and a complete symmetry of the vestibulo-ocular responses was restored in about 3 weeks. It is concluded that a unilateral lesion of the flocculus leads to two separate, but interacting, effects upon vestibulo-ocular responses.

A paradoxical improvement of misreaching in optic ataxia: new evidence for two separate neural systems for visual localization.

We tested a patient (A. T.) with bilateral brain damage to the parietal lobes, whose resulting 'optic ataxia' causes her to make large pointing errors when asked to locate single light emitting diodes presented in her visual field. We report here that, unlike normal individuals, A. T.'s pointing accuracy improved when she was required to wait for 5 s before responding. This counter-intuitive result is interpreted as reflecting the very brief time-scale on which visuomotor control systems in the superior parietal lobe operate. When an immediate response was required, A. T.'s damaged visuomotor system caused her to make large errors; but when a delay was required, a different, more flexible, visuospatial coding system--presumably relatively intact in her brain--came into play, resulting in much more accurate responses. The data are consistent with a dual processing theory whereby motor responses made directly to visual stimuli are guided by a dedicated system in the superior parietal and premotor cortices, while responses to remembered stimuli depend on perceptual processing and may thus crucially involve processing within the temporal neocortex.

Grasping an object: one movement, several components.

The visuomotor transformations for producing a grasping movement imply simultaneous control of different visual mechanisms. The object size, orientation and 3D characteristics have to be encoded for the selection of the appropriate opposition space, within which the opposition forces will be applied on the object surface. These mechanisms also have to combine with those of the transport of the hand to the object location. Finally, biomechanical constraints impose categorical visuomotor decisions for positioning the opposition space according to object changes in size, orientation and spatial location. This paper examines possible interactions between the specialized structures for visuomotor transformation and the internal model that adapts prehension to its goals.