The influence of limb crossing on left tactile extinction

BACKGROUND: Previous research on patients with left tactile extinction has shown that crossing of hands, so that each hand is on the opposite side of the body midline relative to the other, improves detection of stimuli given to the left hand. OBJECTIVES: To study the influence of the spatial position of limbs on left tactile extinction, and its relations with left visual neglect. METHODS: Normal participants and patients with right cerebral hemisphere damage and left tactile extinction were asked to detect single or double light touch stimuli applied to their cheeks, hands, or knees with their arm and legs either in anatomical or in crossed position, increasing the attentional load of the task. RESULTS: In patients with left extinction, limb crossing caused a deterioration in performance for stimuli applied to right body parts, with only a tendency to an improvement in detection for left body parts (only two of 24 patients showed substantial (>20%) improvement in left extinction after limb crossing). After crossing, left limb detections of double stimuli decreased with increasing degrees of visual neglect. CONCLUSIONS: In conditions of high attentional load, limb crossing may impair tactile detection in most patients with left extinction, and particularly in those showing signs of left visual neglect. These results underline the importance of general attentional capacity in determining tactile extinction. Attentional and somatotopic mechanisms of extinction may assume different weights in different patients.     

Lateralization and interhemispheric integration of directed attention]

Directed attention is a function to direct and shift the focus of awareness adequately to behaviorally relevant sensory events. Healthy subjects direct attention evenly to right and left hemispaces. Unilateral spatial neglect is a failure to respond normally to stimuli on the side opposite a cerebral lesion, which is considered to represent a unilateral disruption of directed attention. The established clinical observation that neglect usually occurs after right hemisphere lesions and the results of functional imaging studies suggest the right hemisphere dominance for directed attention. It is hypothesized that the right hemisphere distributes attention to space bilaterally, whereas the left hemisphere distributes attention primarily to right hemispace. However, patients with callosotomy show no apparent neglect with either right or left hand. Ishiai et al. (2001) reported detailed analyses of eye movements when a patient with a callosal infarction bisected lines. Left unilateral spatial neglect may appear, when use of the right hand induces a rightward bias in the attentional control of the left hemisphere and damage to its cingulate gyrus inhibits interhemispheric integration of attention. By contrast, the disconnected but intact right hemisphere may bisect a line accurately by integrating attention to the extents perceived in the left and right visual fields.     

Changes in spatial position of hands modify tactile extinction but not disownership of contralesional hand in two right brain-damaged patients

Somatic misperceptions and misrepresentations, like supernumerary phantom limb and denial of ownership of a given body part, have typically been reported following damage to the right side of the brain. These symptoms typically occur with personal or extrapersonal neglect and extinction of left-sided stimuli, suggesting that all these different symptoms may be linked to the same neural substrate. In the present research, we tested two right brain-damaged (RBD) patients to find out whether changing the position of the hands in space influences tactile extinction and denial of ownership to the same extent. Results showed that manipulation of the spatial position of the hands reduces tactile extinction but leaves denial of ownership of the left hand unaffected. Such a dissociation suggests that delusional misperceptions may be independent from somatic neglect and that representation of hands in space and attribution of ownership are dynamically mapped in at least partly separate neural substrates.     

Changes in Spatial Position of Hands Modify Tactile Extinction but not Disownership of Contralesional Hand in Two Right Brain-Damaged Patients

Abstract

Somatic misperceptions and misrepresentations, like supernumerary phantom limb and denial of ownership of a given body part, have typically been reported following damage to the right side of the brain. These symptoms typically occur with personal or extrapersonal neglect and extinction of left-sided stimuli, suggesting that all these different symptoms may be linked to the same neural substrate. In the present research, we tested two right brain-damaged (RBD) patients to find out whether changing the position of the hands in space influences tactile extinction and denial of ownership to the same extent. Results showed that manipulation of the spatial position of the hands reduces tactile extinction but leaves denial of ownership of the left hand unaffected. Such a dissociation suggests that delusional misperceptions may be independent from somatic neglect and that representation of hands in space and attribution of ownership are dynamically mapped in at least partly separate neural substrates.     

Effects of vestibular rotatory accelerations on covert attentional orienting in vision and touch

Peripheral vestibular organs feed the central nervous system with inputs favoring the correct perception of space during head and body motion. Applying temporal order judgments (TOJs) to pairs of simultaneous or asynchronous stimuli presented in the left and right egocentric space, we evaluated the influence of leftward and rightward vestibular rotatory accelerations given around the vertical head-body axis on covert attentional orienting. In a first experiment, we presented visual stimuli in the left and right hemifield. In a second experiment, tactile stimuli were presented to hands lying on their anatomical side or in a crossed position across the sagittal body midline. In both experiments, stimuli were presented while normal subjects suppressed or did not suppress the vestibulo-ocular response (VOR) evoked by head-body rotation. Independently of VOR suppression, visual and tactile stimuli presented on the side of rotation were judged to precede simultaneous stimuli presented on the side opposite the rotation. When limbs were crossed, attentional facilitatory effects were only observed for stimuli presented to the right hand lying in the left hemispace during leftward rotatory trials with VOR suppression. This result points to spatiotopic rather than somatotopic influences of vestibular inputs, suggesting that cross-modal effects of these inputs on tactile ones operate on a representation of space that is updated following arm crossing. In a third control experiment, we demonstrated that temporal prioritization of stimuli presented on the side of rotation was not determined by response bias linked to spatial compatibility between the directions of rotation and the directional labels used in TOJs (i.e., "left" or "right" first). These findings suggest that during passive rotatory head-body accelerations, covert attention is shifted toward the direction of rotation and the direction of the fast phases of the VOR.     

Neural consequences of somatosensory extinction

There are currently two main interpretations proposing mechanisms underlying tactile extinction: sensory and attention deficit hypotheses. Kinsbourne proposed an opponent processor model to support the attention deficit hypothesis. He insisted that bilateral hemispheres interact reciprocally through contralaterally oriented vectors, and in patients presenting extinction, balance is impaired, causing inattention. From Kinsbourne's point of view, extinction is not caused by sensory disturbance but inattention, therefore even in extinction patients, simultaneous bilateral stimuli should reach the bilateral primary sensory cortices (SI). Using functional magnetic resonance imaging (fMRI), tactile stimuli were administered to both hands of healthy subjects as well as a tactile extinction patient. The patient with tactile extinction extinguished right palm stimuli following simultaneous palm stimulation. During the fMRI study, we gave tactile stimuli to the right palm, the left palm, and simultaneously to both palms. In normal subjects, simultaneous bilateral stimuli activated the bilateral SI and bilateral secondary sensory cortices (SII). In the patient with right tactile extinction, simultaneous bilateral stimuli activated the bilateral SI along with the bilateral SII and right superior parietal lobule. Our study suggests that activation of SI is insufficient to engender an awareness of sensory stimuli. From the view point of Kinsbourne, stimulus driven activity in one hemisphere suppresses activity in the other hemisphere via callosal connections. Our results support the notion that an undamaged superior parietal lobule in the patient with tactile extinction suppresses the damaged parietal lobe function and causes extinction.     

Spatio-temporal prediction modulates the perception of self-produced stimuli.

We investigated why self-produced tactile stimulation is perceived as less intense than the same stimulus produced externally. A tactile stimulus on the palm of the right hand was either externally produced, by a robot or self-produced by the subject. In the conditions in which the tactile stimulus was self-produced, subjects moved the arm of a robot with their left hand to produce the tactile stimulus on their right hand via a second robot. Subjects were asked to rate intensity of the tactile sensation and consistently rated self-produced tactile stimuli as less tickly, intense, and pleasant than externally produced tactile stimuli. Using this robotic setup we were able to manipulate the correspondence between the action of the subjects' left hand and the tactile stimulus on their right hand. First, we parametrically varied the delay between the movement of the left hand and the resultant movement of the tactile stimulus on the right hand. Second, we implemented varying degrees of trajectory perturbation and varied the direction of the tactile stimulus movement as a function of the direction of left-hand movement. The tickliness rating increased significantly with increasing delay and trajectory perturbation. This suggests that self-produced movements attenuate the resultant tactile sensation and that a necessary requirement of this attenuation is that the tactile stimulus and its causal motor command correspond in time and space. We propose that the extent to which self-produced tactile sensation is attenuated (i.e., its tickliness) is proportional to the error between the sensory feedback predicted by an internal forward model of the motor system and the actual sensory feedback produced by the movement.     

Hemispheric specialization for tactile perception opposed by contralateral noise

The aim of this study was to investigate the contribution of each cerebral hemisphere in the categorization of tactile information with and without contralateral tactile noise. Two groups of subjects rated the dissimilarity of paired stimuli varying in shape and texture presented to the left or right hand, with contralateral noise in the other hand for one group only. Analysis of variance on derived structured indices showed a general left hand advantage in the treatment of this haptic information. Moreover, a group by hand interaction revealed that noise-exposed subjects showed better differentiation of tactile information for stimuli presented to the left hand. The data suggest a unidirectional interference by the left hemisphere in the processing of tactile information in noise-free conditions and that quality of interhemispheric transfer of information may modulate cerebral functional lateralization.     

Emotionally negative stimuli can overcome attentional deficits in patients with visuo-spatial hemineglect

Left unilateral spatial neglect resulting from right brain damage is characterized by loss of awareness for stimuli in the contralesional side of space, despite intact visual pathways. We examined using fMRI whether patients with neglect are more likely to consciously detect in the neglected hemifield, emotionally negative complex scenes rather than visually similar neutral pictures and if so, what neural mechanisms mediate this effect. Photographs of emotional and neutral scenes taken from the IAPS were presented in a divided visual field paradigm. As expected, the detection rate for emotional stimuli presented in the neglected field was higher than for neutral ones. Successful detection of emotional scenes as opposed to neutral stimuli in the left visual field (LVF) produced activations in the parahippocampal and anterior cingulate areas in the right hemisphere. Detection of emotional stimuli presented in the intact right visual field (RVF) activated a distributed network of structures in the left hemisphere, including anterior and posterior cingulate cortex, insula, as well as visual striate and extrastriate areas. LVF-RVF contrasts for emotional stimuli revealed activations in right and left attention related prefrontal areas whereas RVF-LVF comparison showed activations in the posterior cingulate and extrastriate visual cortex in the left hemisphere. An additional analysis contrasting detected vs. undetected emotional LVF stimuli showed involvement of left anterior cingulate, right frontal and extrastriate areas. We hypothesize that beneficial role of emotion in overcoming neglect is achieved by activation of frontal and limbic lobe networks, which provide a privileged access of emotional stimuli to attention by top-down modulation of processing in the higher-order extrastriate visual areas. Our results point to the importance of top-down regulatory role of the frontal attentional systems, which might enhance visual activations and lead to greater salience of emotional stimuli for perceptual awareness.     

Neural correlates of conscious and unconscious vision in parietal extinction

Brain areas activated by stimuli in the left visual field of a right parietal patient suffering from left visual extinction were identified using event-related functional magnetic resonance imaging. Left visual field stimuli that were extinguished from awareness still activated the ventral visual cortex, including areas in the damaged right hemisphere. An extinguished face stimulus on the left produced robust category-specific activation of the right fusiform face area. On trials where the left visual stimulus was consciously seen rather than extinguished, greater activity was found in the ventral visual cortex of the damaged hemisphere, and also in frontal and parietal areas of the intact hemisphere. These findings extend recent observations on visual extinction, suggesting distinct neural correlates for conscious and unconscious perception.     

Neural Correlates of Conscious and Unconscious Vision in Parietal Extinction

Brain areas activated by stimuli in the left visual field of a right parietal patient suffering from left visual extinction were identified using event-related functional magnetic resonance imaging. Left visual field stimuli that were extinguished from awareness still activated the ventral visual cortex, including areas in the damaged right hemisphere. An extinguished face stimulus on the left produced robust category-specific activation of the right fusiform face area. On trials where the left visual stimulus was consciously seen rather than extinguished, greater activity was found in the ventral visual cortex of the damaged hemisphere, and also in frontal and parietal areas of the intact hemisphere. These findings extend recent observations on visual extinction, suggesting distinct neural correlates for conscious and unconscious perception.     

The space of senses: impaired crossmodal interactions in a patient with Balint syndrome after bilateral parietal damage

Balint syndrome after bilateral parietal damage involves a severe disturbance of space representation including impaired oculomotor behaviour, optic ataxia, and simultanagnosia. Binding of object features into a unique spatial representation can also be impaired. We report a patient with bilateral parietal lesions and Balint syndrome, showing severe spatial deficits in several visual tasks predominantly affecting the left hemispace. In particular, we tested whether a loss of spatial representation would affect crossmodal interactions between simultaneous visual and tactile events occurring at the same versus different locations. A tactile discrimination task, where spatially congruent or incongruent visual cues were delivered near the patient's hands, was used. Following stimulation of the left hand in the left side of space, we observed visuo-tactile interactions that were not modulated by spatially congruent conditions. In contrast, performance following stimulation of the right hand in the right side of space was affected in a spatially selective manner--facilitated for congruent stimuli and slowed for incongruent stimuli. To dissociate effects on somatotopic and spatiotopic coordinates, we crossed the patient's hands during unimodal tactile discriminations. Tactile performance of the left hand improved when it was positioned in the right hemispace, whereas placing the right hand in left space produced no significant changes, suggesting that left-sided tactile inputs are coded with respect to a combination of limb- and trunk-centred coordinates. These data converge with recent findings in animals and healthy humans to indicate a critical role of the posterior parietal cortex in multimodal spatial integration, and in the fusion of different coordinates into a unified representation of space.     

Changes of muscle excitability of the hand contralateral to a task performing one: a transcranial magnetic stimulation study]

It used to be considered that unilateral movements of distal limb parts are associated only with contralateral motor cortical activity. Recent neuroimaging studies, however, suggest that the motor cortex ipsilateral to a task-performing hand is also activated, and that motor patterns in one hand affect the degree of the activity of the ipsilateral motor cortex. If so, muscles of the hand contralateral to a task-performing one may change those excitability depending on types of tasks. We studied eight subjects who performed three different finger tasks by one hand: (a) pinch, (b) sequential finger opposition, and (c) tactile discrimination. Transcranial magnetic stimulation was delivered by a figure eight coil over the hemisphere ipsilateral to a task-performing hand. Motor evoked potentials and background electromyographic activities were recorded from the opponens pollicis muscle contralateral to the stimulated hemisphere. On average, the motor evoked potentials were larger during tactile discrimination task than those at rest in either hand (p < 0.01). Background electromyographic activities in the left hand increased significantly during right hand tactile discrimination task (p < 0.01), whilst those in the right hand did not change during the left hand performance (p > 0.05). These findings suggest the followings: (1) the hand muscle contralateral to a task performing one changes its excitability depending on types of tasks; and (2) increment of excitability of the left hand muscle associated with right hand tactile discrimination is greater than that of the right hand one in association with the same task by the left hand, thus supporting the idea that there is a functional asymmetry between the right and left motor cortex in respect of motor performance.     

The cerebellum is involved in predicting the sensory consequences of action.

We used H2(15)O PET to examine neural responses to parametrically varied degrees of discrepancy between the predicted and actual sensory consequences of movement. Subjects used their right hand to move a robotic arm. The motion of this robotic arm determined the position of a second foam-tipped robotic arm, which made contact with the subject's left palm. Using this robotic interface, computer controlled delays were introduced between the movement of the right hand and the tactile stimulation on the left. Activity in the right lateral cerebellar cortex showed a positive correlation with delay. These results suggest the cerebellum is involved in signalling the sensory discrepancy between the predicted and actual sensory consequences of movements.     

Hemispheric differences in dichaptic scanning of verbal and spatial material by adult males and females

An experimental design based on the combination of dichaptic presentation associated to the Posner’s paradigm was adopted to investigate laterality effects for verbal and spatial non-linguistic stimuli in male and female adult normal subjects. In a grapheme similarity judgment task based on “Name Identity” a right hand/left hemisphere advantage was found. Conversely, laterality effects were neither observed when the task involved “Perceptually Identical” or “Different” letter pairs. In a further experiment, the same methodology was adopted to verify hemispheric effects with spatial non-linguistic material, and a significant advantage for the left hand/right hemisphere was observed. Contrary to many previous studies, no gender or gender × task effects have been detected in both experiments. The present results suggest the existence, also in the tactile domain, of a direct link between input type and the linguistic or non-linguistic processing to which the two hemispheres are devoted. The overall pattern of data seriously hampers Witelson’s [Cortex 10 (1974) 3] original hypothesis that letter stimuli presented in the tactile modality are primarily processed as spatial stimuli, and are therefore dependent on the right hemisphere functioning.     

Temporal pattern of odor administration alters hemispheric processing in humans

Evidence from a variety of sensory modalities has suggested that the left hemisphere may be 'tuned' to process more rapidly changing stimuli than the right and some have suggested that this difference forms the foundation of the functional dichotomy often drawn between the two hemispheres. Odors may be thought to engage these same temporally dependent processes as portions of an odor mixture may come to be transduced into a phasic series of neural events. Using brain electrical activity, we show that the temporal sequence of the odor alters the pattern of brain electrical activity. Estimates of the source localization for this activity indicate that rapidly changing odors, like sounds, visual and tactile stimuli, show increased activity in the left hemisphere.     

Visuo-tactile integration in personal space

Integration of information across sensory modalities is enhanced when stimuli in both modalities are in the same location. This "spatial rule" of multisensory integration has been primarily studied in humans by comparing stimuli located either in the same versus opposite side of the body midline or in peripersonal versus extrapersonal space, both of which involve large, categorical differences in spatial location. Here we used psychophysics and ERPs to investigate visuo-tactile integration in personal space (i.e., on the skin surface). We used the mirror box technique to manipulate the congruence of visual and tactile information about which finger on either the right or left hand had been touched. We observed clear compatibility effects for both visual and tactile judgments of which finger on the left hand had been touched. No such effects, however, were found for judgments about the right hand. ERP data showed a similar pattern. Amplitude of the vertex P200 potential was enhanced and that of the N2 was reduced for congruent visuo-tactile events on the left, but not the right, hand. Similarly, a later positivity over posterior parietal cortices (P300) showed contralateral enhancement for congruent visuo-tactile events on both the left and right hands. These results provide clear evidence for spatial constraints on visuo-tactile integration defined in personal space and also reveal clear lateralization of these effects. Furthermore, these results link these "ultraprecise" spatial constraints to processing in the right posterior parietal cortex.     

Hemispheric function in disorganized type schizophrenia: performance on the quality extinction test.

We assessed hemisphere function in right-handed male chronic, disorganized type schizophrenic patients (N = 60, age range 18-45 years) using the Quality Extinction Test (QET), in comparison to 20 right-handed male healthy controls in the same age range. The QET analysis discriminated between the disorganized schizophrenic patients and the controls. QET results indicated that chronic schizophrenic patients were less sensitive to tactile stimuli in both hands as compared to controls. Furthermore, the sensitivity to tactile stimuli of the left hand was less than that of the right hand in the schizophrenic patients. In contrast, in the normal controls the sensitivity was similar in both hands. These results indicate possible right hemisphere dysfunction together with disturbance in interhemispheric transmission through the corpus callosum in chronic, disorganized type schizophrenic patients.     

Asymmetric Influence of Egocentric Representation onto Allocentric Perception

Objects in the visual world can be represented in both egocentric and allocentric coordinates. Previous studies have found that allocentric representation can affect the accuracy of spatial judgment relative to an egocentric frame, but not vice versa. Here we asked whether egocentric representation influenced the processing speed of allocentric perception. We measured the manual reaction time of human subjects in a position discrimination task in which the behavioral response purely relied on the target's allocentric location, independent of its egocentric position. We used two conditions of stimulus location: the compatible condition-allocentric left and egocentric left or allocentric right and egocentric right; the incompatible condition-allocentric left and egocentric right or allocentric right and egocentric left. We found that egocentric representation markedly influenced allocentric perception in three ways. First, in a given egocentric location, allocentric perception was significantly faster in the compatible condition than in the incompatible condition. Second, as the target became more eccentric in the visual field, the speed of allocentric perception gradually slowed down in the incompatible condition but remained unchanged in the compatible condition. Third, egocentric-allocentric incompatibility slowed allocentric perception more in the left egocentric side than the right egocentric side. These results cannot be explained by interhemispheric visuomotor transformation and stimulus-response compatibility theory. Our findings indicate that each hemisphere preferentially processes and integrates the contralateral egocentric and allocentric spatial information, and the right hemisphere receives more ipsilateral egocentric inputs than left hemisphere does.     

Effects of preferred hand and sex on the perception of tactile simultaneity

The effects of handedness, sex and the influence of hand placement in extrapersonal space on temporal information processing was investigated by measuring thresholds for perceiving the simultaneity of pairs of tactile stimuli. Simultaneity thresholds of preferred right handed and left handed university students with left hemisphere speech representation were compared using unimanual and bimanual stimulation at three hand placements (midline, lateral and crossed). In unimanual conditions two fingers of one hand were stimulated (single hemisphere), whereas in the bimanual conditions one finger of each hand was stimulated (cross hemispheres). Bimanual minus unimanual thresholds provided an estimate of interhemisphere transmission time (IHTT) regardless of hand placement. The effects of hemispace varied with the type of stimulation. With unimanual stimulation, overall thresholds were longer at the midline placement, however, with bimanual stimulation, thresholds were longer when the hands were spatially separated (crossed and/or uncrossed). Left handers' IHTTs were 8 ms faster than those of right handers. IHTTs in males were faster than females with hands placed in lateral (by 10.8 ms) or crossed (by 9.8 ms) but not midline positions. It was concluded that the cerebral hemispheres are equally capable of discriminating temporal intervals, but that the left hemisphere predominates when there is uncertainty about location of stimulation.