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The cingulum is a core component of the limbic lobe and part of the circuit that was described by Papez where environmental experiences become endowed with emotional awareness. Recent techniques for the study of cerebral connectivity have updated this fasciculus' morphology and led to the acknowledgment that its involvement in superior functions goes far beyond emotion processing. Long and robust, the cingulum is a long association fasciculus with terminations in all cerebral lobes. These observations plead for a pivotal rethinking of its role in the human brain and lead to the conclusion that to merely consider it as the main fasciculus of the limbic system was actually a reductionism. This paper summarizes the key facts regarding why the cingulum is now perceived as a primary interconnecting apparatus in the medial aspect of the cerebral hemisphere.

We present an extensive investigation (24 experiments) of a new case of prosopagnosia following right unilateral damage, GG, with the aim of addressing two classical issues: (1) Can a visual recognition impairment truly be specific to faces? (2) What is the nature of acquired prosopagnosia? We show that GG recognizes nonface objects perfectly and quickly, even when it requires fine-grained analysis to individualize these objects. He is also capable of perceiving objects and faces as integrated wholes, as indicated by normal Navon effect, 3D-figures perception and perception of Mooney and Arcimboldo face stimuli. However, the patient could not perceive individual faces holistically, showing no inversion, composite, or whole-part advantage effects for faces. We conclude that an occipito-temporal right hemisphere lesion may lead to a specific impairment of holistic perception of individual items, a function that appears critical for normal face recognition but not for object recognition.

Callosal disconnection can reveal asymmetrical contributions of the two brain hemispheres to praxis. In this paper, we revisit a study of a patient with callosal disconnection (Goldenberg et al., 2001, Neuropsychologia, 39:1432-1443), who perfectly imitated meaningless gestures when imitation was controlled only by the left hemisphere, but was severely impaired when the right hemisphere was in charge of motor control. We decomposed the gestures into a set of geometric variables that were to be reproduced, such as the orientation of the hand and the position of contact between the hand and the face. Whereas orientation of the hand in extrinsic coordinates was replicated correctly by both hemispheres, only the left hemisphere reproduced correctly the position of contact between the hand and the face. This goal-dissociation as well as several partial perseveration errors speak against the hypothesis of a direct route from perception to motor replication of gestures, as interruption of a direct route would probably impair all the features of the gesture. We speculate that incorrect coordination between the reproductions of multiple goals may be the core deficit underlying callosal apraxia.

We investigated whether or not there is a relationship between hand preference for grasping and hemispheric dominance for language—and how each of these is related to other traditional measures of handedness. To do this we asked right- and left-handed participants to put together two different sets of 3D puzzles made out of big or very small LEGO © pieces. Participants were also given two self-reported handedness questionnaires, as well as tests of grip force and finger tapping speed. A language lateralization (dichotic listening) test was also administered. We found a positive correlation between hand use for precision grasping and language lateralization (i.e. the more participants used their right hand for grasping the small LEGO© pieces, the more language was lateralized to the left hemisphere). In addition, we identified two populations of left-handers according to their grasping performance: ‘left-right-handers’, who behaved exactly like right-handers; and ‘left-left-handers’ whose performance was the mirror image of that of right-handers. Finally, we found an increase in right-hand use when right-handers and ‘left-righthanders’ had to pick up the small LEGO© pieces. We discuss our results in relation to recent notions of lefthemisphere specialization for visually guided actions and its relationship with the evolution of language.

Goldenberg [Goldenberg, G. (1996). Defective imitation of gestures in patients with damage in the left or right hemisphere. Journal of Neurology, Neurosurgery, and Psychiatry, 61, 176-180] proposed that the vulnerability of the imitation of meaningless gestures to right or left brain damage depends on the body parts that are involved in the gestures. Whereas imitation of hand postures was disturbed only in patients with left brain damage, imitation of finger postures was affected to similar degrees in left and right brain damage. Subsequent studies confirmed the selective vulnerability of hand postures to LBD but failed to replicate the severe disturbance of finger postures in RBD. In contrast to Goldenberg's studies, these studies excluded RBD patients with neglect. The present investigation aimed to explore the relationship between spatial neglect and imitation of finger postures in RBD patients. Presence and severity of spatial neglect and accuracy of imitation of hand and finger postures were tested in 50 RBD patients. Disturbance of imitation was much more severe for finger than hand postures and was tightly correlated with severity of neglect. The number of errors was higher for fingers which from the patient's perspective were located on the left side of the examiner's demonstrating hand but this spatial bias was not sufficient to explain all errors. Possible causes for non-lateralized errors could be a general narrowing of the focus of attention and reduced capacity for processing of visual information which have been postulated to be regular companions of the lateral displacement of attention in spatial neglect.

Although most cases of acquired prosopagnosia are accompanied by bilateral brain lesions, a number also arise following right unilateral lesions. The prevailing consensus is that right hemisphere damage disrupts the configural apprehension of faces, which in turn forces a reliance on part-based processing. Here we describe a patient who following right hemisphere damage is not only unable to apprehend the configural aspects of faces, but is also unable to apprehend their component parts when these are presented within a whole, upright face. Intriguingly, the patient is able to apprehend face parts when these are presented in isolation, within inverted faces, or in unfamiliar, scrambled arrangements. Furthermore, the patient can make use of configural information to detect local changes in non-face stimuli. The findings uncover a hitherto unreported form of impairment following right unilateral damage, and raise questions about the role of the left hemisphere in processing local information.

Event-related potential (ERP) studies have shown that sensitivity to individual faces emerges as early as approximately 160ms in the human occipitotemporal cortex (N170). Here we tested whether this effect generalizes across changes in viewpoint. We recorded ERPs during an unfamiliar individual face adaptation paradigm. Participants were presented first with an adapting face ( approximately 3000ms) rotated 30 degrees in depth, followed by a second face (200ms) in a frontal view of either the same or a different identity. The N170 amplitude at right occipitotemporal sites to the second stimulus was reduced for repeated as compared to different faces. A bilateral adaptation effect emerged after 250ms following stimulus onset. These observations indicate that individual face representations activated as early as 160ms after stimulus onset in the right hemisphere show a substantial degree of generalization across viewpoints.

We have studied the configuration of the cortico-subcortical language networks within the right hemisphere (RH) in nine left-handers, being operated on while awake for a cerebral glioma. Intraoperatively, language was mapped using cortico-subcortical electrostimulation, to avoid permanent deficit. In frontal regions, cortical stimulation elicited articulatory disorders (ventral premotor cortex), anomia (dorsal premotor cortex), speech arrest (pars opercularis), and semantic paraphasia (dorsolateral prefrontal cortex). Insular stimulation generated dysarthria, parietal stimulation phonemic paraphasias, and temporal stimulation semantic paraphasias. Subcortically, the superior longitudinal fasciculus (inducing phonological disturbances when stimulated), inferior occipito-frontal fasciculus (eliciting semantic disturbances during stimulation), subcallosal fasciculus (generating control disturbances when stimulated), and common final pathway (inducing articulatory disorders during stimulation) were identified. These cortical and subcortical structures were preserved, avoiding permanent aphasia, despite a transient immediate postoperative language worsening. Both intraoperative results and postsurgical transitory dysphasia support the major role of the RH in language in left-handers, and provide new insights into the anatomo-functional cortico-subcortical organization of the language networks in the RH-suggesting a "mirror" configuration in comparison to the left hemisphere.

During visually guided foraging birds tend to select certain types of food from a mixed diet. This selectivity is ecologically relevant. During scanning for food birds spot the surroundings mainly with the monocular lateral visual field of the one or other eye and then control pecking with their small binocular frontal visual field. As the visual systems of the avian left and right brain hemisphere are supposed to work largely independently in the short term, the problem arises of how the avian brain handles a task that requires coordinated activity of the left and right brain hemisphere for efficient processing. Here we report that chicks exhibit strong selective feeding when both of the brain hemispheres are involved. With the left or right hemisphere alone selectivity is reduced or completely absent. Our findings reveal a marked qualitative difference between unilateral and bilateral processing. They highlight an important but so far unexplored selection pressure for the evolution of hemispheric cooperation.

Previous research suggests that the right hemisphere (RH) may contribute uniquely to the processing of metaphoric language. However, most studies have focused on familiar metaphoric expressions. The present study used the divided visual field paradigm to examine the role of the right cerebral hemisphere in processing novel metaphoric expressions taken from poetry. In two experiments, participants were presented with four types of word pairs, literal, conventional metaphoric and novel metaphoric expressions and unrelated word pairs, and asked to perform a semantic judgment task. Two different SOAs: 400 and 1100 ms were used. The results of both experiments showed that responses to LVF/RH presented target words were more accurate and faster than responses to RVF/LH target words for novel metaphoric expressions, but not for other types of word pairs. These results support previous research indicating that during word recognition, the RH activates a broader range of related meanings than the LH, including novel, nonsalient meanings. The findings thus suggest that the RH may be critically involved in at least one important component of novel metaphor comprehension, i.e., the integration of the individual meanings of two seemingly unrelated concepts into a meaningful metaphoric expression.

Auditory temporal processing was investigated in individuals with acquired aphasia using a task in which they were asked to detect brief silent gaps inserted between noise segments modeled after formants in speech. To examine within-channel gap detection, gaps of 10, 20, 40, and 80ms duration were inserted between an initial segment (IS) and a trailing segment (TS) centered at the same frequency (1kHz). In a between-channel gap detection condition, gaps of 20, 40, 80, and 100ms duration were inserted between an IS that differed in frequency (4kHz) from the TS (1kHz). The effect of gap onset timing was examined in both conditions by systematically varying the duration of the IS by 10, 20, or 40ms. A combined analysis revealed that for both conditions and all gap and IS durations, individuals with aphasia produced fewer correct responses than age-matched neurologically intact controls. Separate condition analyses revealed that when noise segments were centered at the same frequency, individuals with aphasia demonstrated poorer accuracy in detecting 40 and 80ms gaps relative to normal controls (p0.001). When gaps were inserted between noise segments differing in frequency, on average, aphasic subjects performed less accurately at durations of 40, 80 and 100ms (p0.025). Detection in both groups decreased with smaller IS durations. The difficulties with gap detection observed in the aphasic group suggest the existence of fundamental problems in processing the temporal form or microstructure of sounds characterized by rapidly changing onset dynamics.

Cerebral asymmetries are thought to be associated with increased hemispheric specialization of function. We investigated cerebral petalias, the protrusion of one cerebral hemisphere relative to the other, and their relationship to lateralized behavior in capuchin monkeys (Cebus apella). Magnetic resonance images of the brain and behavioral data on a coordinated bimanual task were obtained from 13 capuchins. While a significant population-level left-frontal petalia was found, this was not related to handedness. The role of the morphologically asymmetric frontal cortex in capuchins is unclear, but may reflect developmental gradients or directional selection for various behavioral functions, such as extractive foraging or social group complexity.

Affective (emotional) prosody is a neuropsychological function that encompasses non-verbal aspects of language that are necessary for recognizing and conveying emotions in communication, whereas non-affective (linguistic) prosody indicates whether the sentence is a question, an order or a statement. Considerable evidence points to a dominant role for the right hemisphere in both aspects of prosodic function. However, it has yet to be established whether separate parts of the right hemisphere are involved in processing different kinds of emotional intonation. The aim of this study was to answer this question. In addition, the issue of sex differences in the ability to understand prosody was considered. Fifty-two patients with damage to frontal, temporo-parietal or subcortical (basal) parts of the right hemisphere and 26 controls were tested for their ability to assess prosody information in normal (well-formed) sentences and in pseudo-sentences. General impairment of prosody processing was seen in all patient groups but the effect of damage was more apparent for emotional rather than linguistic prosody. Interestingly, appreciation of emotional prosody appeared to depend on the type of emotional expression and the location of the brain lesion. The patients with frontal damage were mostly impaired in comprehension of happy intonations; those with temporo-parietal damage in assessment of sad intonations, while subcortical lesions mostly affected comprehension of angry intonations. Differential effects of lesion location on the performance of men and women were also observed. Frontal lesions were more detrimental to women, whereas subcortical lesions led to stronger impairment in men. This suggests sex differences in brain organization of prosodic functions.

We used a computerized delayed-reaching task with a simple reaction time (RT) to investigate the visuo-motor and spatio-temporal performance of right brain-damaged (RBD) patients with unilateral spatial neglect (USN). Fifty-three RBD patients (22 with and 31 without USN) and 25 controls performed the tasks. We recorded the following data: the first RT (RT-1), which is thought to reflect the detection of the target position (the perceptual factor); the second RT (RT-2), which represents the initiation of reaching (the motor initiation aspect of premotor factors); the movement time (MT), which is hypothesized to reflect the "pure" motor component of the task. RBD patients with both USN and hemianopia demonstrated significantly longer RTs towards the left than towards the right for both the RT-1 and the RT-2. Among the RBD patients without hemianopia, the laterality index (left side/right side) of the RT-1 in those with USN was significantly greater than in those without USN or the controls. Among the three groups, there were no significant differences between the laterality indices of either the RT-2s or the MTs. These results suggest that the impairment of leftward movement in RBD patients with USN might be caused primarily by a perceptual impairment rather than an impairment in motor initiation, and is certainly not a "pure" motor impairment.

We used fMRl to study the neural correlates of crossmodal, visual-tactile extinction in a single case (patient GK). GK has chronic extinction after a lesion centred on right inferior parietal cortex, and has previously been investigated extensively in purely visual fMRI studies [e.g. Rees, G., Wojciulik, E., Clarke, K., Husain, M., Frith, C., & Driver, J. (2000). Unconscious activation of visual cortex in the damaged right hemisphere of a parietal patient with extinction. Brain, 123(Pt 8), 1624-1633; Rees, G., Kreiman, G., & Koch, C. (2002). Neural correlates of consciousness in humans. Nature Reviews Neuroscience, 3(4), 261-270]. With concurrent stimulation of the right visual field plus left index finger, GK showed crossmodal extinction of left touch on approximately half of such trials here (reflecting impaired sensitivity, i.e. lowered d-prime), albeit becoming aware of left touch on the other half. fMRI revealed activation of contralateral primary somatosensory cortex on crossmodal trials when touch was extinguished from awareness, suggesting unconscious residual processing there. When GK became aware of the left touch, additional activation was found in surviving right parietal cortex, and in frontal regions; moreover, functional coupling was enhanced with a region of frontal cortex implicated in awareness by previous work. Finally, on trials where crossmodal extinction arose, surviving right parietal cortex showed stronger functional coupling with the left visual and right somatosensory regions driven by the competing stimuli, indicating that crossmodal extinction arises when inputs to separate modalities interact competitively via multimodal cortex. (c) 2006 Elsevier Ltd. All rights reserved.

The coordination of sensorimotor tasks involving both visual half-fields requires exchange of information between the brain hemispheres. So far, this interhemispheric information transfer has never been investigated under conditions where the two hemispheres receive different visual inputs and each hemisphere performs on a different task. The present study asked whether such conditions affect the transfer of information between the hemispheres, and if so, at which processing stage. We addressed these issues by pairing a face comparison task with a visual discrimination task (I-task) designed to interfere with the interhemispheric information transfer required for face comparison. One version of the I-task (experiment 1) required discrimination of the faces of John Lennon and Yoko Ono; the other version (experiment 2) required discrimination between the names 'JOHN' and 'YOKO'. Thus, the two I-task versions overlapped at early visual processing stages where visual feature analysis is carried out, but differed at later processing stages where words or faces are represented as objects. We found that both I-task versions disrupted the interhemispheric information transfer for the face comparison task. This indicates that when both hemispheres are occupied by separate tasks, interhemispheric communication is less efficient. In addition, our results suggest that the hemispheres exchange sensory information already at a rather early visual processing stage. Hence, visual feature analysis in one hemisphere is probably informed about feature analysis in the other hemisphere and may also be modulated by it.

The present study describes in detail, for the first time, a case of failure with multiplication procedures in a right hemisphere damaged patient (PN). A careful, step-by-step, error analysis made possible to show that an important portion of PN's errors could be better explained as spatial in nature and specifically related to the demands of a multi-digit multiplication. These errors can be distinguished from other types of errors, including those, expected after a right hemisphere lesion, determined by a generic inability to deal with spatial material, or from other deficits, like neglect, affecting cognitive capacities across the board. The best explanation for PN's problems is that he might have difficulties in relying on a visuo-spatial store containing a layout representation specific to multiplication. As a consequence, while knowing what, when and how to carry out the various steps, PN does not know where. What he may thus lack is a spatial schema of multiplication. Such schema is thought to help normal calculators in overcoming working memory demands of complex calculation by representing the information of where exactly each sub-step should be placed.

Despite the extensive clinical and neuropsychological evidence for major differences in language function between the left and right hemispheres, it has proven difficult to demonstrate reliable differences in lexical processing. It is particularly surprising that frequency has been found to have no differential effects upon word recognition in the hemispheres, in view of the evidence that this variable has significant effects upon lexical processing. It is possible, however, that any interaction between word frequency and hemispheric processes is too subtle or too complex to be revealed by the simple dichotomous studies previously attempted. This study manipulated word frequency across seven levels in an effort to provide a fine-grained insight into the relationship between word frequency and lexical processing in the hemispheres. The results revealed a clear additivity between visual field and word frequency across all levels of frequency, for both RT and errors. Such findings create difficulties for the interpretation of studies of lateralized lexical access in general, and are particularly hard to reconcile with lateral asymmetries revealed in other aspects of lexical processing. It is proposed that a resolution of this problem may depend upon the development of theories that emphasise the notion of laterally distributed word representations, rather than discrete lexical structures on each side of the brain.

Joke comprehension deficits in patients with right hemisphere (RH) damage raise the question of the role of the intact RH in understanding jokes. One suggestion is that semantic, or meaning, activations are different in the RH and LH, and RH meanings are particularly important for joke comprehension. To assess whether hypothesized differences in semantic activation in the two hemispheres were relevant to joke comprehension, we recorded event-related brain potentials (ERPs) as healthy adults read laterally presented “punch words” to one-line jokes and nonjoke controls. Jokes presented to the RVF/LH elicited larger amplitude N400 than the nonjoke endings; when presented to the LVF/RH, the joke and nonjoke endings elicited N400s of equal amplitude. This finding suggests that semantic activations in the two hemispheres do differ, with RH semantic activation facilitating joke comprehension. © 2004 Elsevier Ltd. All rights reserved.

The attentional blink (AB) refers to a decrement in detecting the occurrence of a probe item if it closely follows a previous target item in rapid serial visual presentation (RSVP). In the present study we presented target and probe stimuli in two parallel RSVP streams, one in each visual field, in order to address the question of whether the AB might differ between the cerebral hemispheres. The characteristic AB, with reduced detection of the probe at post-target Lags 2-5, but no such deficit at Lag 1 (Lag 1 sparing), was observed when target and probe were both in the right visual field. When they were both presented in the left visual field the AB was attenuated. When the target and probe were in opposite visual fields, probe detection was again reduced when it was in the left visual field, and there was no Lag 1 sparing. The left-visual-field advantage in performing the AB task may reflect a general right-hemispheric specialization for attentional processing.

Auditory extinction was examined in six patients with right hemisphere lesions, using an auditory double simultaneous stimulation task using letters spoken in male and female voices. Patients had to detect where the stimuli were located, and identify either the letter or the voice (the task-relevant dimension). Auditory extinction was greatest when the two stimuli were the same on the task-relevant dimension, paralleling previous studies of visual extinction. Furthermore, we found that errors of omission were much less frequent in the contralesional field if the patients were not required to report both identity and the location of stimuli. These results are consistent with the notion that auditory extinction may be due in part to a failure to bind together identity and location.

In patients with right brain damage (RBD) or left brain damage (LBD) and healthy subjects, tactile and three basic gustatory stimuli (sour, salty, bitter) were applied to the left or right hemitongues or to both hemitongues simultaneously. Tactile stimuli were detected and localized by verbal report, whereas gustatory stimuli were identified by pointing to the corresponding name on cards bearing the names of the three tastes. In the tactile test, 9 of 18 RBD patients showed extinction of left hemitongue stimuli, whereas the remaining RBD patients, 9 LBD patients and 14 healthy subjects detected virtually all stimuli in all conditions. In the gustatory test, healthy subjects outperformed the two brain damaged groups which nevertheless responded well above chance and did not differ from one another. Unexpectedly, the nine RBD patients with left hemitongue tactile extinction showed no gustatory extinction, since performance did not differ significantly between the two hemitongues on both unilateral and bilateral stimulations. To account for these findings, some evidence suggests that the tongue representation is bilateral in both modalities, but predominantly ipsilateral in the gustatory modality and predominantly contralateral in the tactile modality. The RBD patients with left hemitongue tactile extinction were those with more marked symptoms of left-sided extinction in the visual and auditory modalities, making it likely that their brain damage was also responsible for left lingual tactile extinction. The absence of left gustatory extinction in those patients can be attributed to the predominant channelling of left hemitongue taste inputs into the intact left hemisphere.

Five people lacking the corpus callosum (two callosotomized, three with agenesis of the corpus callosum) and neurologically normal subjects were shown vertical lines that appeared instantaneously between pairs of rectangles in one or other visual field. When one of the rectangles flashed prior to the presentation of the line, and the line was in the same visual field, all subjects perceived the line as spreading from the flashed rectangle to the other. Normal subjects and one of the callosotomized subjects showed a slight but significant right visual-field advantage, perhaps reflecting a left-hemispheric superiority in processing rapid temporal events. The illusion was also induced when the line and the flash were in opposite visual fields in one of the callosotomized, one of the acallosal subjects, and about half of the normal subjects, implying interhemispheric integration even in the absence of the corpus callosum.

It is widely assumed that the local and global levels of hierarchical stimuli are processed more efficiently in the left and right cerebral hemispheres, respectively. However, corresponding effects were not observed under all circumstances. In ERP studies, they occurred more often with centrally presented stimuli than with laterally presented ones, whereas reaction-time studies revealed that a response conflict between the levels is relevant. The present study examines which of these two factors is more important by presenting conflicting and non-conflicting stimuli to the left or right visual field and recording ERPs as well as collecting behavioral data. If a central stimulus position is crucial, then no effects should show up. Contrary to this prediction, the expected hemispheric differences were observed in the behavioral data as well as in the later occurring (N2 and P3) ERP amplitudes. However, in all variables, the effects were more pronounced for conflicting stimuli. The results suggest that response conflicts are more important for obtaining hemispheric differences in global/local processing than a central stimulus presentation. This is interpreted in the way that hemispheric differences vary with respect to the stimulus representation that is needed to select a proper response.

A speeded lexical decision task was used to investigate word-category deficits in patients suffering from lesions in the right hemisphere and in neurological controls without cortical lesion. In all patients from one group (n = 12), the right frontal lobe was affected causing a left-sided hemiparesis. In the second group (n = 6), lesions primarily affected areas in the right inferior temporo-occipital lobes. Patients with motor deficits due to lesions in the spinal cord or in the periphery served as neurological controls (n = 9). Processing of three categories of words was investigated: verbs referring to actions (action verbs (acVs)); nouns with strong visual associations (visually-related nouns (viNs)); and nouns with both strong action and visual associations (bimodal nouns (biNs)). Stimulus categories were matched for word length and normalized lexical frequency. Error scores revealed a significant word category by patient group interaction. Patients with lesions in the right frontal lobe showed most severe deficits in processing action verbs, whereas those with lesions in their right temporo-occipital areas showed most severe deficits in processing visually-related nouns. Neurological controls did not show any differences between word categories. The double dissociation of the processing impairments seen in frontal versus temporo-occipital patients demonstrates that specific word-category deficits can arise from lesions in the right non-dominant hemisphere. An account for these results in terms of distributed neuronal systems representing words is offered.

Converging evidence suggests that temporal representations of brief durations are derived subcortically. We tested split-brain patient JW in order to investigate whether these representations project bilaterally or unilaterally to cortex. Using visual stimuli to signal time intervals, JW was asked to compare the duration of a pair of standard stimuli that were presented bilaterally with a comparison stimulus that was presented to either the left or right visual field. Assuming the hand of response is controlled by the contralateral cerebral hemisphere, a hand by visual field interaction was predicted if the representation of stimulus duration was restricted to the cerebral hemisphere receiving the lateralized stimulus. However, we failed to observe this interaction for two different ranges of stimulus durations, both in the hundred (Experiment 2) to hundreds (Experiment 1) of milliseconds range. Instead, there was a consistent right hemisphere advantage in task performance. When the task then required a discrimination based on the physical size of the stimuli rather than their duration, an interaction between response hand and visual field was obtained (Experiment 3). Taken together, these results suggest that (1) even though the comparison stimulus was presented unilaterally, the representation of its duration was available to both cerebral hemispheres, and (2) a right hemisphere advantage in psychophysical tasks requiring the comparison of successive stimuli is observed for temporal and non-temporal judgments.

We examined the electrophysiological correlates of left-sided tactile extinction in a patient with right-hemisphere damage. Computer-controlled punctate touch was presented to the left, right or both index fingers in an unpredictable sequence. The patient reported his conscious tactile percept ("left", "right" or "both"). He showed extinction on 75% of bilateral trials, reporting only right stimulation for these. Somatosensory evoked potentials for unilateral stimulation showed early components over contralateral somatosensory areas (P60 and N110) for either hand. In contrast to the results observed for age-matched controls, the patient's P60 was smaller in amplitude for left-hand touch over the right hemisphere than for right-hand touch over the intact hemisphere. Bilateral trials with extinction revealed residual P60 and N110 components over the right hemisphere in response to the extinguished left touch. These results demonstrate residual unconscious somatosensory processing of extinguished touch. They also suggest that tactile extinction can be caused by attenuation rather than elimination of somatosensory responses in the damaged hemisphere, with an underlying deficit even on unilateral trials.

We administered 24 min of unilateral vibrotactile stimulation to the left or right ventral forearm of experimental participants. A separate control group did not receive stimulation but completed all experimental tasks. Participants who received right-side stimulation persisted more in attempting to solve insoluble puzzles and also made more positive judgments about an emotionally neutral film compared to participants who received left-side stimulation. The performance of the control group was intermediate to that of the stimulation groups, but was significantly different only from the right stimulation group. The absence of a significant left-side stimulation effect is not readily interpretable because floor effects may have limited the attenuating effects of left-side stimulation on persistence in problem solving. Emotional biases in cognition and performance resulting from unilateral vibrotactile stimulation can be explained with reference to activation of structures involved in the processing of emotion within the contralateral cerebral hemisphere, and by the laterality of emotion specified in the Valence Hypothesis. These results support the contention that similar biases elicited by unilateral muscle contractions result from their sensory consequences.

Implicit, high level processing of extinguished objects has often been described in the visual modality. In the tactile domain, however, research on this topic is meagre and it is still uncertain whether processing of tactually presented stimuli can be affected by the same attentional disorders as visual stimuli. In this paper we describe a patient, ENM, with visual neglect and light touch extinction who, in a naming task of objects presented in the tactile modality, simultaneously to both hands, showed extinction for left hand objects. He was, nevertheless, able to make above chance Same/Different judgements on the two stimuli. We also tested two neurologically intact subjects who performed the test wearing a ski-glove on the left hand to impair the recognition of left hand objects. In these subjects, Same/Different judgements were at chance level when recognition rate was as low as that found in patient ENM. This happened when either the objects, although sharing the same name were different in shape (conditions Same-Different) or when the two objects were different with respect to the category name but were actually physically similar (conditions Different-Similar). However, when the objects were either identical or completely different, i.e., in a condition where judgement could be based simply on the physical analysis of the object shape (condition Same Identical and Different Dissimilar), their Same/Different judgements were above chance, despite the tactual deficit. Our conclusion was that patient ENM showed implicit recognition of left hand objects, at least in the Same Different and in the Different-Similar conditions, whereas, in the same conditions, normal subjects with an artificial sensory impairment did not. Our results also show that Same/Different judgements may be, in some conditions, less demanding than naming tasks, as suggested by Farah et al. Furthermore, patient ENM performed the test both with uncrossed and crossed hands. We found that extinction always affected the hand contralateral to the brain damage, although there was a tendency for a decrement of the ipsilesional hand performance in the crossed condition. We discuss these findings with reference to the most recent theories on the existence of a body centered spatial frame of reference.

An unselected group of right hemisphere, semi-acute stroke patients ( n =30) was run on a laboratory test of naturalistic action production and was found to commit errors of action at a higher rate than what was previously reported for recovering head injury patients [Schwartz et al., Naturalistic action impairment in closed head injury. Neuropsychology , 1997, 8, 59–72]. There were strong similarities in how these two patient groups responded to variations in task demands and in the pattern of errors they produced. Hemispatial biases were evident in the errors of right hemisphere patients with neglect but not those without neglect; and neglect patients also many errors that were unrelated to the spatial layout. We argue that a non-specific resource limitation — which might translate as reduced arousal or effort — is central to the breakdown of naturalistic action production after brain damage, and right hemisphere patients are especially vulnerable to this resource limitation and its behavioral consequences.

Joanette and Goulet ( Neuropsychologia , 1986, 24, 875–879) reported that right-handed patients suffering from a right-hemisphere stroke were impaired on semantic but not on orthographic criteria when they were submitted to a word fluency task. The interpretation of this dissociation was that the right hemisphere would subtend a particular contribution to the semantic component of word processing in right handers. The goal of the research reported here was to challenge this dissociation. In contrast with the first study, the production criteria were presented randomly and their level of productivity was a controlled factor. The semantic-orthographic dissociation was not replicated under these conditions.

Thirty-one right-hemisphere lesioned (RHL) patients, 11 left-hemisphere lesioned patients (LHL) and 10 normal controls (NC) bisected lines in three spatial location and four directional cuing conditions. The error direction and error size were analyzed as separate and combined variables. Seventy-seven percent of RHL patients and 45% of LHL patients made abnormally large errors in line bisection. Right-hemisphere lesioned patients were more sensitive to spatial location and directional cuing than NC subjects. In contrast, LHL patients were less sensitive to either condition than NCs. The error direction and error size emerged as dissociable components of line bisection. Right-hemisphere lesioned patients and NC subjects bisected lines consistently to one side of the true center. Left-hemisphere lesioned patients bisected lines equally often on both sides of the true center. Both RHL and LHL patients made larger absolute bisection errors than NC subjects, but the RHL patient's errors were larger than those of the LHL patients. We propose that the greater sensitivity of RHL patients to spatial location and directional cues and the directional consistency of their bisection errors represent contributions of the intact left cerebral hemisphere to line bisection. In contrast, the LHL patient's unrestrained ability to orient to both ends of the line reflects a contribution of the intact right cerebral hemisphere to line bisection. The failure of both groups to accurately bisect lines reflects a common visuospatial processing deficit that is more pronounced following RHLs than LHLs.

Children with pre- or perinatal injury to right hemisphere (RH) brain regions show impairment of spatial integrative functions similar to that observed among adults with comparable injury. Unlike adults, children show considerable improvement with development on a range of spatial construction tasks which require spatial integration. Such gains could reflect true recovery of spatial integrative abilities. Alternatively, the improvement could be more limited in scope, reflecting the development of compensatory strategies which are task specific and allow the children to circumvent, rather than overcome, their primary spatial disorders. The studies presented here examined this distinction within the context of drawing tasks in which the child was first asked to draw a house and then an impossible house. The impossible house task was designed to examine the extent to which children rely on graphic formulas in generating organized drawings. The results showed that while all of the children with RH injury make considerable progress in free drawing into the school age period, they are very reliant on the use of graphic formulas. When given a task which requires them to alter their drawings, they did not change the spatial configuration of the depicted object. Rather they found alternate ways to render the object ‘impossible’.

The case of a professional musician with a right temporal lobe epilepsy is presented. Whilst playing an organ concert (John Stanley's Voluntary VIII, Op. 5), he suffered a complex partial seizure. The recorded concert performance (with the seizure) was analysed and compared with other available exercise records and with the composition. The musical analysis of the seizure-induced variations reveals that at the beginning of the seizure, the left hand started to become unprecise in time and deviated from the score, whereas the right hand remained faultless at this time. With increasing duration of the seizure discharge, the dissociation of both hands from the score increased but the right hand compensated for the errors of the left hand in a musically meaningful way, i.e. with the aim to compensate for the seizure-induced errors of the left hand. The case illustrates untroubled musical judgement during epileptic activity in the right temporal lobe at the beginning of the seizure. Whereas the temporal formation of the performance was markedly impaired, the ability of improvisation—in the sense of a `perfect musical solution' to errors of the left hand—remained intact.

The early position that prosopagnosia is predominantly associated with right hemisphere (RH) injury was challenged by the finding that in practically all cases that come to autopsy pathological data point to bilateral damage. Yet the rejection of the RH hypothesis may have been too hasty. We report three prosopagnosic patients in whom MRI and CT documented a lesion confined to the right occipito-temporal areas and PET confirmed that hypometabolism involved the RH only. A review of the literature brought out 27 cases with neuroimaging evidence that prosopagnosia was associated with RH damage plus four cases with surgical evidence. It remains, however, that the inability to recognize familiar faces is a rare disorder, not manifested by the majority of patients with right temporo-occipital injury. We submit that right-handers differ in the degree of their RH specialization in processing faces and that in only a minority of them is it so marked that it cannot be compensated for by the healthy left hemisphere.

We have studied multi-joint arm movements in a patient with right cerebellar damage. Our patient's kinetic tremor was reduced in the absence of visual feedback, but increased when performing a concurrent cognitive task. We suggest that her kinetic tremor reflects use of a slow cortical feedback circuit for error-corrections during movement. Analysing the spatial relation between hand aperture and hand transport in prehensile movements, we found preserved strategic, trial-to-trial coordination, but impaired reactive, within-trial coordination. We conclude that the proprioceptive representations provided by the normal cerebellum play an important role in coordinating multi-joint movement.

It is controversial whether a stimulus projected within 1 to 3 degrees from the boundary between the right and left hemiretina is transmitted to only one cerebral hemisphere or to both cerebral hemispheres. In order to resolve this issue, letter- and word-stimuli were presented for 200 msec with a new type of tachistoscope, called the fundus tachistoscope, in and about the central retina, (i.e. fovea, 1.2 degrees in horizontal diameter) of the right eyes of two commissurotomized subjects (N.G. and A.A.). During stimulus presentation the subjects were attempting to fixate a fixation target. The fundus tachistoscope combined with image analysis of the fundus enables us to measure the position of the stimulus on the retina, relative to the foveal center, as well as whether or not the eye moved during stimulus presentation. The results indicate that the region of the right (temporal) hemiretina represented by both hemispheres in letter processing, if it exists, was estimated as less than 0.6 degrees from the foveal center. The two subjects frequently (27% in N.G. and 46% in A.A.) fixated the fixation target eccentrically, i.e. with a retinal point other than the foveal center, during fixation, namely stimulus presentation. Their eccentric fixations were small with magnitude almost all falling between 1.35 degrees right and 1.25 degrees left of the foveal center. It is therefore recommended that letter-stimuli be presented at least 2.0 degrees from the foveal center in ordinary tachistoscopic studies of cerebral hemispheric differences. Eye movements, which varied in 0.11 degrees and 1.43 degrees horizontally, occurred in about 8% of all the trials during fixation. On the average of the two subjects, the eye movements caused or worsened eccentric fixation in only about one third of the trials, and corrected eccentric fixation in about two thirds of the trials.

A brain activation study using SPECT and 133Xe in a deep dysphasic patient with left temporal lesion is presented. The activation paradigm consisted of a passive listening to foreign language as baseline, a phoneme monitoring condition and a semantic word monitoring condition. The specific activation of the right middle temporal cortex observed in the semantic condition is congruent with the hypothesis of a compensatory role of the right hemisphere in processing concrete words. This case illustrates the interest of functional imaging for a better understanding of neural mechanisms of functional recovery after brain injury.

A 2-[18F]-Fluoro-2-Deoxy- D -Glucose ([18F]FDG) and positron emission tomography (PET) study was performed in the acute and chronic phase of stroke in one patient with unilateral neglect due to a right hemispheric lesion. In the acute phase, severe neglect, as well as hypometabolism in both the right and in the left unaffected cerebral hemisphere, was demonstrated. At follow-up evaluation the patient showed an almost complete recovery fromunilateral neglect. This was associated with a return of left hemisphere metabolism to normal values and partial metabolic recovery in the right hemisphere, where frontal and parietal areas remained functionally impaired. Another patient with an extensive right cerebral ischaemic lesion on CT and severe unilateral neglect was studied by PET in chronic phase. A severe metabolic depression in the left unaffected hemisphere and in the right cerebral areas spared by the lesion, was found. These data suggest that the remission of unilateral neglect might be associated to a functional metabolic recovery in both the undamaged left hemisphere and the unaffected regions of the right hemisphere.

Frontal lobe and basal ganglia lesions have been associated with similar cognitive impairments, although their specialized roles in behavior are likely to be different. We examined whether these structures mediate distinctive or overlapping aspects of a complex behavioral process that has been associated with both neural sites, i.e. cognitive flexibility. Patients with focal ischemic lesions to the frontal lobe and basal ganglia were compared on two forms of cognitive flexibility: (1) shifting response set (i.e. reactive flexibility), and (2) producing a diversity of ideas (i.e. spontaneous flexibility). Results indicated that frontal lobe and basal ganglia damage each caused a similar degree of impairment in reactive flexibility, both groups performing at a significantly lower level than posterior cortical lesion and normal comparison groups. However, frontal lobe damage markedly disturbed spontaneous flexibility, while performance after basal ganglia lesion was significantly higher and comparable to posterior cortical lesions. Findings suggest that the frontal lobe and basal ganglia participate differently in the neural substrate of cognitive flexibility. The frontal lobe appears to mediate spontaneous flexibility. The production of diverse ideas may require direct cortical-cortical interactions by the frontal lobe in order to access knowledge systems with novel strategies that transcend the most common semantic linkages. In contrast the corticostriate system appears to mediate reactive fexibility, as the frontal lobe, basal ganglia and their interconnections are required for its operation.

The rightward line bisection errors made by patients with visuospatial neglect can be explained as due to a spatially misdirected response, which would be predicted on either of two accounts. An alternative view, however, is that such patients actually misperceive the left half of a horizontal line as being shorter than the right half. We have tested this possibility directly in three neglect patients, by giving them prebisected lines: they were found to judge a central transection mark as lying nearer to the left end of the lines. We were also able to test one of the patients on a series of size comparisons using computer-generated patterns. She was found to judge horizontal lines as shorter in the left half of visual space than in the right. This was also true for comparisons of the areas of nonsense figures. However she did not make such constant errors when comparing the lengths of vertical lines. It is suggested that an attentional deficit in left hemispace may result in the underestimation of horizontal extent. This would act in combination with misdirected reaching to determine the magnitude of line bisection errors.

Ideomotor limb apraxia, commonly defined as a disorder of skilled, purposeful movement, is characterized by spatiotemporal deficits during a variety of actions. These deficits have been attributed to damage to, or impaired retrieval of, stored representations of learned actions, especially object-related movements. However, such deficits might also arise from impaired visuomotor transformation mechanisms that operate in parallel to or downstream from mechanisms for storage of action representations. These transformation processes convert extrinsic visual information into intrinsic neural commands appropriate for the desired motion. These processes are a key part of the movement planning process and performance errors due to inadequate transformations have been shown to increase with the dynamic complexity of the movement. This hypothesis predicts that apraxic patients should show planning deficits when reaching to visual targets, especially when the coordination and/or dynamic requirements of the task increase. Three groups (18 healthy controls, 9 non-apraxic and 9 apraxic left hemisphere damaged patients) performed reaching movements to visual targets that varied in the degree of interjoint coordination required. Relative to the other two groups, apraxic patients made larger initial direction errors and showed higher variability during their movements, especially when reaching to the target with the highest intersegmental coordination requirement. These problems were associated with poor coordination of shoulder and elbow torques early in the movement, consistent with poor movement planning. These findings suggest that the requirement to transform extrinsic visual information into intrinsic motor commands impedes the ability to accurately plan a visually targeted movement in ideomotor limb apraxia.

Following stroke, a patient may fail to report touch administered by another person but claim that she feels touch when it is self-administered. We investigated three explanations for self-touch enhancement: (1) proprioceptive information from the administering hand, (2) attentional modulation, and (3) temporal expectation. Tactile sensation was assessed with vision precluded, and with the affected hand positioned in the left and right hemispace. In four of six experiments, the somatic rubber hand paradigm was used: the Examiner administered stimulation to the patient's affected left hand while guiding the patient's right hand to administer synchronous stimulation to a prosthetic hand. Even though the patient's two hands were not in contact, patients detected the same number of stimulations as when they touched their own hand directly (self-administered condition). Moreover, there was no decline in rates of detection when potentially informative movements of the administering hand were restricted. This demonstrates that patients feel rather than infer stimulation under conditions of self-touch. When patients received stimulation to the affected hand in the opposite hemispace to the hand administering touch to the prosthetic hand, all but one showed self-touch enhancement. Thus, neither proprioceptive information nor attentional modulation at the spatial region of the administering hand provided a sufficient explanation for self-touch enhancement. A follow-up experiment indicated an important role for temporal expectation: a delay, between the patient's stimulation of the prosthetic hand and the Examiner's stimulation of the patient's affected hand, eliminated the self-touch enhancement effect.

Twenty-seven patients with right cerebral infarcts resulting in left-sided neglect in a line cancellation task were also tested for line cancellation when the lines were confined to the left half of sheets. All but one patient (whose neglect in the former condition was minimal) were still impaired in the latter condition, a finding that indicates that hyperattraction to test stimuli in the unneglected (right) hemispace was not the crucial factor preventing them from completing the standard version of the cancellation task, although it may have played some role. The results are consistent with the conclusion that hyperattention to stimuli in the unneglected hemifield, and possibly to stimuli in the unneglected hemispace, together with other factors, could account for the patients' impairment in line cancellation.

Eight subjects performed physical and nominal letter-matching tasks for pairs of letters presented in the left, right, or central visual fields, using a unimanual two-choice response-time paradigm. Latencies were manipulated by using a cued response procedure, and speed-sensitivity and speed-bias functions were calculated separately for each of the peripheral visual field by response hand conditions. Hemispheric contributions to these tasks were investigated by looking for evidence of exclusive specialization and callosal transfer. The results suggest that both hemispheres are able to perform the tasks and that they do so using similar processing strategies, but that they differ in the response-choice stage of the nominal identity task.

Volpe et al. (Nature 282, 722, 1979 [19]) described an experimental study of four patients with parietal tumours who were able to judge whether two simultaneous stimuli were identical or different, even when they were unable to name the stimulus contralateral to their brain injury. We report the case of another patient, E.M., in whom we have investigated this phenomenon further. E.M. had undergone a right temporal lobectomy to prevent recurrent seizures. She could correctly name photographs of objects presented in isolation to either the left or right visual field, at 150 msec exposure (although she was impaired for single objects on the left at 10 msec exposures). She was able to judge correctly whether two simultaneous objects on the left and right had the same or different names, even though she was often unable to name the object on the left. These judgements remained above chance when same-name pairs of stimuli showed the same object but seen from two different viewpoints, or even when they showed visually dissimilar exemplars of the same name category. This implies that the patient based her same-different judgements on categorical information about the pair of objects, even though she was often unable to name the contralateral object.

Ninety-four epileptic patients who had undergone the carotid amytal test were evaluated in order to determine whether sex affects the pattern of hemispheric reorganization following cerebral injury. Our data suggest that the period during which hemispheric reorganization for speech can occur differs between the sexes. Females have a relatively short window, with reorganization most likely during the first year of life. The length of this window is more difficult to define for males but may extend until puberty. The period during which a shift to left-handedness can occur does not appear to be affected by gender. Here, both males and females seem to show a relatively short window.

This study investigated the effect of exemplar dominance on semantic priming in the left and right visual fields for words that are members of the same category, but not strongly associated. A low proportion of related primes was employed in lexical decision and word pronunciation tasks to assess the automatic activation of word meanings in each cerebral hemisphere. Priming was reliably obtained only in the LVF/right hemisphere. In addition, this effect did not vary with category dominance of the prime:equivalent LVF priming was observed for ROBIN-CROW (high dominant) and DUCK-CROW (low dominant) pairs. These findings support the view that a broader range of related meanings is activated during word recognition in the right, than in the left, hemisphere.

Intentionality, or Theory of Mind, is the ability to explain and predict the behaviour of others by attributing to them intentions and mental states and is hypothesised to be one of several social cognitive mechanisms which have impacted upon brain size evolution. Though the brain activity associated with processing this type of information has been studied extensively, the neuroanatomical correlates of these abilities, e.g. whether subjects who perform better have greater volume of associated brain regions, remain to be investigated. Because social abilities of this type appear to have evolved relatively recently, and because the prefrontal cortex (PFC) was the last brain region to develop both phylogenetically and ontogenetically, we hypothesised a relationship between PFC volume and intentional competence. To test this, we estimated the volume of four regional prefrontal subfields in each cerebral hemisphere, in 40 healthy adult humans by applying stereological methods on T1-weighted magnetic resonance images. Our results reveal a significant linear relationship between intentionality score and volume of orbital PFC (p = 0.01). Since this region is known to be involved in the processing of social information our findings support the hypothesis that brain size evolution is, at least in part, the result of social cognitive mechanisms supporting social cohesion.

Split fovea theory proposes that when the eyes are fixated within a written word, visual information about the letters falling to the left of fixation is projected initially to the right cerebral hemisphere while visual information about the letters falling to the right of fixation is projected to the left cerebral hemisphere. The two parts of the word must be re-united before the word can be recognised. Bilateral projection theory proposes instead that visual information is projected simultaneously to both hemispheres provided that it falls within the fovea (defined as the central 2–3°). On this more traditional account, no interhemispheric transfer would be required in order to read a word presented within the fovea. We review the evidence in support of split fovea theory and consider some of the objections that have been raised. We argue that a split fovea affects the reading of words at fixation, something that must be recognised and accounted for by cognitive, computational and neural models of reading.