Writing with the right hemisphere

We studied writing abilities in a strongly right-handed man following a massive stroke that resulted in virtually complete destruction of the language-dominant left hemisphere. Writing was characterized by sensitivity to lexical-semantic variables (i.e., word frequency, imageability, and part of speech), semantic errors in writing to dictation and written naming, total inability to use the nonlexical phonological spelling route, and agrammatism in spontaneous writing. The reliance on a lexical-semantic strategy in spelling, semantic errors, and impaired phonology and syntax were all highly consistent with the general characteristics of right hemisphere language, as revealed by studies of split-brain patients and adults with dominant hemispherectomy. In addition, this pattern of writing closely resembled the syndrome of deep agraphia. These observations provide strong support for the hypothesis that deep agraphia reflects right hemisphere writing.     

Effect of hemispherectomy in infantile hemiplegics

Four patients with infantile hemiplegia who had undergone hemispherectomy were examined. both left (LH) and right (RH) hemidecorticates obtained borderline defective scores on IQ tests. Patients with isolated right hemispheres were found not to be aphasic or apraxic. Visuospatial and constructional capacities were compromised in all patients. Individuals with an isolated right hemisphere smiled and gestured more than did those with an isolated right hemisphere. EEG and CT scan data showed that the remaining hemisphere was not normal in these patients. Therefore, inferences regarding the functional plasticity of the remaining hemisphere must be made with caution.     

When only the right hemisphere is left: studies in language and communication

An adult of above normal intelligence, BL, underwent left hemispherectomy at age five, and subsequently graduated from college and has been regularly employed. Using standardized neuropsychological instruments, previous extensive testing had revealed optimal performance for a hemispherectomized subject. To probe communicative abilities in greater detail, and to examine current questions about linguistic superiority of the left hemisphere and "crowding" of right hemisphere functions, 12 additional protocols were administered. BL performed at normal or above on nearly all protocols. However, performance on production of phonemically complex words was effortful, and deficits were seen on two tests requiring comprehension of linguistic contrasts in prosody (Linguistic Prosody Test) and syntax (the Active-Passive Test). These findings support previous claims of reduced ability in specific, circumscribed linguistic functions in the left hemispherectomized person, and lead to suggestions for further testing of communicative competence in individuals with a single intact hemisphere.     

Social environment elicits lateralized navigational paths in two populations of typically developing children

The current study provides the first evidence of human lateralized navigation of a social space within a naturalistic environment. We employed a quantitative, observational approach and report on a detailed set of nearly 700 independent navigational routes from two separate child populations consisting of over 300 typically developing children, aged five to fourteen years. The navigational path was considered across the sagittal plane (left, right) around three distinct target types (peer, adult and object). Both child populations expressed a significant bias for choosing a rightward navigational path around a human target (e.g. peer, adult) and no lateral preference for navigation around fixed, inanimate objects. A rightward navigational path provides an advantage for the left visual field and the right hemisphere, facilitating both the production and perception of social-emotion stimuli. The findings are consistent with evidence from studies of non-human animal species demonstrating that the social environment elicits predictable lateralized behavior, and support an early evolutionary delineation of functional processing by the two hemispheres.     

How normal is grammatical development in the right hemisphere following hemispherectomy? The root infinitive stage and beyond

We examined the morphosyntax of eight left hemispherectomized children at two different stages and compared it to MLU-matched normals. We found that the language of the hemispherectomies paralleled that of their MLU matches with respect to the specific morphosyntactic characteristics of each stage. Our findings provide strong evidence for the presence of functional categories in all early grammars and demonstrate that grammatical development, regardless of its neural substrate, is highly constrained by UG and follows a narrowly determined course. We discuss our findings within a neurobiological framework in which etiology defines the integrity of the remaining hemisphere, which in turn, determines its potential for linguistic reorganization and/or acquisition.     

Language after hemispherectomy

We studied the spoken language of 49 children who had undergone hemispherectomy as part of the UCLA Pediatric Epilepsy Surgery Research Program and analyzed, among a number of clinical factors, the relation between acquired vs developmental pathology and spoken language outcomes. In this paper we will briefly review the results of our study and attempt to explain (1) why "the early" is not always better, (2) why so many right hemispherectomies fail to develop language, and (3) why some left hemispherectomized children develop remarkably good language despite removal of the "language" hemisphere. This account will rest on the proposed model of brain maturation and progressive lateralization.     

The angular gyrus in developmental dyslexia: task-specific differences in functional connectivity within posterior cortex

Converging evidence from neuroimaging studies of developmental dyslexia reveals dysfunction at posterior brain regions centered in and around the angular gyrus in the left hemisphere. We examined functional connectivity (covariance) between the angular gyrus and related occipital and temporal lobe sites, across a series of print tasks that systematically varied demands on phonological assembly. Results indicate that for dyslexic readers a disruption in functional connectivity in the language-dominant left hemisphere is confined to those tasks that make explicit demands on assembly. In contrast, on print tasks that do not require phonological assembly, functional connectivity is strong for both dyslexic and nonimpaired readers. The findings support the view that neurobiological anomalies in developmental dyslexia are largely confined to the phonological-processing domain. In addition, the findings suggest that right-hemisphere posterior regions serve a compensatory role in mediating phonological performance in dyslexic readers.     

Time sharing and dichotic listening asymmetry in normal and learning-disabled children

Time-sharing and dichotic listening techniques were used to examine cerebral lateralization for language function in 48 normal and 48 learning-disabled children. All subjects were matched according to age, sex, and handedness. An analysis of results indicated that both nornal and learning-disabled children demonstrated left hemisphere lateralization of language function on the time-sharing and dichotic listening tasks. However, no developmental trends were evident for either group. Differences observed in the performance of the normal and learning-disabled children may relate to how each group utilizes “verbal strategies” and processes simultaneous information in the left hemisphere. The results strongly question the notion that attributes learning disabilities to incomplete or delayed language lateralization and lend support to the notion that cerebral lateralization is not a developmental phenomena.     

Right-hemisphere reading in a case of developmental deep dyslexia

The right-hemisphere hypothesis of deep dyslexia has received support from functional imaging studies of acquired deep dyslexia following damage to the left cerebral hemisphere, but no imaging studies of cases of developmental deep dyslexia, in which brain damage is not suspected, have been reported. In this paper, we report the first evidence of right hyperactivation in an adult case of developmental deep dyslexia. Hyperactivation was observed in the right inferior frontal cortex during functional magnetic resonance imaging (fMRI) of the oral reading of imageable content words and nonwords to which imageable lexical responses were frequently made. No evidence of right hyperactivation was observed in the oral reading of function words, nor during the naming of imageable words in response to pictured objects. The results reveal strategic and selective use of right-hemisphere functions for particular types of written stimuli. We propose that children with developmental deep dyslexia compensate for their lack of phonological skills by accessing right-hemisphere imageable associations that provide a mnemonic for linking written forms to spoken names.     

Right-hemisphere reading in a case of developmental deep dyslexia

The right-hemisphere hypothesis of deep dyslexia has received support from functional imaging studies of acquired deep dyslexia following damage to the left cerebral hemisphere, but no imaging studies of cases of developmental deep dyslexia, in which brain damage is not suspected, have been reported. In this paper, we report the first evidence of right hyperactivation in an adult case of developmental deep dyslexia. Hyperactivation was observed in the right inferior frontal cortex during functional magnetic resonance imaging (fMRI) of the oral reading of imageable content words and nonwords to which imageable lexical responses were frequently made. No evidence of right hyperactivation was observed in the oral reading of function words, nor during the naming of imageable words in response to pictured objects. The results reveal strategic and selective use of right-hemisphere functions for particular types of written stimuli. We propose that children with developmental deep dyslexia compensate for their lack of phonological skills by accessing right-hemisphere imageable associations that provide a mnemonic for linking written forms to spoken names.     

Writing induces a right hemisphere linguistic advantage in dysphonetic dyslexic children: implications for attention and capacity models of laterality

An experiment demonstrated a complete hemispheric processing reversal in 10 male, dysphonetic dyslexic children that occurred during a dichotic listening test of their verbal working memory. Requiring a written response to dichotic digits produced a right hemisphere/left ear superiority in the dysphonetic dyslexics whereas normal subjects and other dyslexics maintained a left hemisphere/right ear advantage. This reversal was unaffected by changes in task difficulty. A second experiment assessed the influence on producing the reversal of concurrent manual interference with left hemisphere verbal processing (responding orally vs. manually) and selective right hemisphere priming (Forward Writing vs. Backward Writing). The dysphonetic children reverted to a strong left hemisphere superiority when recalling the dichotic digits orally. Backward writing produced no ear advantage in either direction. The findings suggest that dysphonetic dyslexia may be related to (1) left hemisphere processing demands that exceed capacity, (2) easily activated right hemisphere processing strategies and (3) failure to coordinate linguistic processing interhemispherically. The results supported a novel hybrid conceptualization of dyslexia consisting of a synthesis of selective activation, and dual processor-limited capacity, theories.     

Real-life spatial skills, handedness, and family history of handedness

According to , pronounced left hemisphere lateralization for language abilities in women, as in female absolute right-handers, limits their right hemisphere capacity and spatial abilities. This study examines the degree of handedness and the family history of non-right-handedness with respect to real-life spatial abilities in women. Twenty-four women had, first, to learn a new route and, second, to orient themselves within a labyrinth. In the former task, the number of errors and completion time were evaluated; in the latter task, degree of error for orienting was recorded. The results show that, contrary to Annett's prediction, right-handers with and without a family history of non-right-handedness did not differ on these measures. In addition, and unexpectedly, absolute right-handers were found to surpass non-absolute ones in the spatial orientation task. These findings do not support Annett's hypothesis and are discussed in relation to functional cerebral organization.     

Encouraging pointing with the right hand,but not the left hand,gives right-handed 3-year-olds a linguistic advantage

Previous research has shown a strong positive association between right-handed gesturing and vocabulary development. However, the causal nature of this relationship remains unclear. In the current study, we tested whether gesturing with the right hand enhances linguistic processing in the left hemisphere, which is contralateral to the right hand. We manipulated the gesture hand children used in pointing tasks to test whether it would affect their performance. In either a linguistic task (verb learning) or a non-linguistic control task (memory), 131 typically developing right-handed 3-year-olds were encouraged to use either their right hand or left hand to respond. While encouraging children to use a specific hand to indicate their responses had no effect on memory performance, encouraging children to use the right hand to respond, compared to the left hand, significantly improved their verb learning performance. This study is the first to show that manipulating the hand with which children are encouraged to gesture gives them a linguistic advantage. Language lateralization in healthy right-handed children typically involves a dominant left hemisphere. Producing right-handed gestures may therefore lead to increased activation in the left hemisphere which may, in turn, facilitate forming and accessing lexical representations. It is important to note that this study manipulated gesture handedness among right-handers and does therefore not support the practice of encouraging children to become right-handed in manual activities.

Research Highlights

• Right-handed 3-year-olds were instructed to point to indicate their answers exclusively with their right or left hand in either a memory or verb learning task.
• Right-handed pointing was associated with improved verb generalization performance, but not improved memory performance.
• Thus, gesturing with the right hand, compared to the left hand, gives right-handed 3-year-olds an advantage in a linguistic but not a non-linguistic task.
• Right-handed pointing might lead to increased activation in the left hemisphere and facilitate forming and accessing lexical representations.

     

Does the right hemisphere take over after damage to Broca's area? the Barlow case of 1877 and its history

In 1877 Thomas Barlow, a London physician, published a remarkable case of functional recovery of speech following brain damage. It involved a 10-year-old boy who had lost his speech, regained it, and lost it again before he died from a disorder that affected his heart and produced embolisms that subsequently affected other organs, including his brain. Examination of the boy's brain revealed two focal regions of softening; one that affected Broca's area and the left facial-motor area, and another, which occurred weeks later, in the homologous regions of the right hemisphere. Although Barlow was most concerned with motor deficits, others at the turn of the century began to cite this case as strong evidence that the corresponding region of the right hemisphere can take over speech functions for Broca's area on the left. Whether this case really provides good support for functional takeover or vicariation theory is critically evaluated in the light of contemporary research, including PET scan studies involving damage to Broca's speech region.     

The neural bases of complex tool use in humans

The behaviors involved in complex human tool use cut across boundaries traditionally drawn between social, cognitive, perceptual and motor processes. Longstanding neuropsychological evidence suggests a distinction between brain systems responsible for representing: (1) semantic knowledge about familiar tools and their uses, and (2) the acquired skills necessary for performing these actions. Contemporary findings in functional neuroimaging support and refine this distinction by revealing the distributed neural systems that support these processes and the conditions under which they interact. Together, these findings indicate that behaviors associated with complex tool use arise from functionally specialized networks involving temporal, parietal and frontal areas within the left cerebral hemisphere.     

Action observation network in childhood: a comparative fMRI study with adults

Very little is known about the action observation network and the mirror neuron system (AON/MNS) in children and its age‐related properties compared with those observed in adults. In the present fMRI study we explored the activation of areas belonging to the AON/MNS in children and adults during observation of complex hand‐grasping actions, as compared to observation of simple grasping acts executed with the left and the right hand, seen from a first person perspective. The results indicate that during the action observation tasks in children there was activation of a cortical network similar to that found in adults, including the premotor cortex, the posterior part of the inferior frontal gyrus and the posterior parietal lobe. However, the activation in children was more widespread and showed a higher inter‐subject variability compared with adults. Furthermore, the activated network seems more lateralized to the left hemisphere in adults and more bilateral in children, with a linear growth of lateralization index as a function of age. Finally, in children the activation in the anterior intraparietal cortex (AIP) of each hemisphere was higher during observation of the contralateral hand (hand identity effect) and during the observation of complex actions relative to simple grasping acts, confirming the role of AIP for action‐related hand identity previously described in adults. These results support the assumption that structure and size of action representations are sensitive to mechanisms of development and show physiological plasticity. These properties of the AON/MNS could constitute a powerful tool for spontaneous reorganization and recovery of motor deficits after brain injury in children and in adults, as well as for specific rehabilitation programmes.     

Neurology of affective prosody and its functional-anatomic organization in right hemisphere

Unlike the aphasic syndromes, the organization of affective prosody in brain has remained controversial because affective-prosodic deficits may occur after left or right brain damage. However, different patterns of deficits are observed following left and right brain damage that suggest affective prosody is a dominant and lateralized function of the right hemisphere. Using the Aprosodia Battery, which was developed to differentiate left and right hemisphere patterns of affective-prosodic deficits, functional-anatomic evidence is presented in patients with focal ischemic strokes to support the concepts that (1) affective prosody is a dominant and lateralized function of the right hemisphere, (2) the intrahemispheric organization of affective prosody in the right hemisphere, with the partial exception of Repetition, is analogous to the organization of propositional language in the left hemisphere and (3) the aprosodic syndromes are cortically based as part of evolutionary adaptations underlying human language and communication.     

Hemispheric differences in case processing

Morphosyntactic capacities of normal brain hemispheres were compared in lexical decision studies involving centrally and laterally presented Serbo-Croatian nouns in different cases. Cases are distinguished by different suffixes and syntactic roles. Experiment 1 confirmed and extended previous findings of the nominative superiority effect: words in the nominative case were processed faster and more accurately than words in other three cases, and nonwords in the nominative case led to more false positive reactions than nonwords in other cases. In Experiment 2 this effect was replicated for right visual field stimuli: nominatives had faster reaction times and smaller error rates than accusatives, and the reversed pattern was found for nonwords. For left visual field stimuli, only the word error analysis found the nominative superior, while the other three analyses (word reaction times, nonword reaction times, and nonword error rates) showed no significant case effect. Word familiarity had an equally strong effect in both hemispheres. The results suggest that centrally presented stimuli are processed by the left hemisphere, that laterally presented stimuli are processed by the initially receiving hemisphere, and that the right hemisphere has a frequency-sensitive lexicon. Reduced right-hemisphere sensitivity for case differences may be due to different lexicon structure or the absence of appropriate morphological or syntactic mechanisms.     

Spatial analysis after perinatal stroke: patterns of neglect and exploration in extra-personal space

This study was conducted to determine whether school-aged children who had experienced a perinatal stroke demonstrate evidence of persistent spatial neglect, and if such neglect was specific to the visual domain or was more generalized. Two studies were carried out. In the first, 38 children with either left hemisphere (LH) or right hemisphere (RH) damage and 50 age-matched controls were given visual cancellation tasks varying in two factors: target stimuli and stimulus array. In the second study, tactile neglect was evaluated in 41 children with LH or RH damage and 72 age-matched controls using a blindfolded manual exploration task. On the visual cancellation task, LH subjects omitted more target stimuli on the right, but also on the left, compared with controls. Children with RH lesions also produced a larger number of omissions on both the left and right sides than controls, but with poorer performance on the left. On the manual exploration task, LH children required significantly longer times to locate the target on both sides of the board than did controls. RH children had significantly prolonged search times on the left side, but not on the right, compared with controls. In both tasks, LH subjects employed unsystematic search strategies more often than both control and RH children. The search strategy of RH children also tended to be erratic when compared to controls, but only in the random arrays of the visual cancellation tasks; structure of the target stimuli improved their organization. These results demonstrate that children with early LH brain damage display bilateral difficulties in visual and tactile modalities; a pattern that is in contrast to that seen in adults with LH damage. This may result from disorganized search strategies or other subtle spatial or attentional deficits. Results of performance of RH children suggests the presence of contralateral neglect in both the visual and tactile modalities; a finding that is similar to the neglect in adult stroke patients with RH lesions. The fact that deficits in spatial attention and organizational strategies are present after very early focal damage to either the LH or the RH broadens our understanding of the differences in functional lateralization between the immature and mature brain. These results also add to evidence for limitations to plasticity in the developing brain. Our findings may have therapeutic and rehabilitative implications for the management of children with early focal brain lesions.