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.     

Selective loss of complex-pitch or speech discrimination after unilateral lesion

Twenty-eight right-handed patients who suffered a single cerebrovascular accident in the distribution of either the left or right middle cerebral artery were tested on their ability to discriminate complex-pitch and speech stimuli presented dichotically. Whereas the left hemisphere lesion group was impaired in dichotic speech but not in dichotic complex-pitch discrimination, the right hemisphere lesion group was impaired in dichotic complex pitch but not in dichotic speech discrimination. Complex-pitch phenomena may provide a useful model for the study of auditory function in the nondominant hemisphere.     

Multiple component agraphia in a patient with atypical cerebral dominance: an error analysis

A 52-year-old man with atypical cerebral dominance (left-handed for writing but mixed handedness for other tasks) suffered an extensive right hemisphere stroke, resulting in a combination of deficits that has not been previously reported. There were profound visual constructive and visual perceptual disturbances and a spatial agraphia, which were consistent with a nondominant hemisphere lesion. There was also a severe apraxic agraphia, which is typically associated with a dominant hemisphere lesion, but no other signs of dominant hemisphere dysfunction such as linguistic disturbance or limb-motor apraxia were present. This case serves to highlight the functional and anatomical relationship between handwriting and other forms of praxis; the various sources of error in letter formation; the need to be specific in labeling and describing agraphias ; and the role of a detailed analysis of writing errors in delineating the neuropsychological processes involved in handwriting.     

Bilateral capacity for speech sound processing in auditory comprehension: evidence from Wada procedures

Data from lesion studies suggest that the ability to perceive speech sounds, as measured by auditory comprehension tasks, is supported by temporal lobe systems in both the left and right hemisphere. For example, patients with left temporal lobe damage and auditory comprehension deficits (i.e., Wernicke's aphasics), nonetheless comprehend isolated words better than one would expect if their speech perception system had been largely destroyed (70-80% accuracy). Further, when comprehension fails in such patients their errors are more often semantically-based, than-phonemically based. The question addressed by the present study is whether this ability of the right hemisphere to process speech sounds is a result of plastic reorganization following chronic left hemisphere damage, or whether the ability exists in undamaged language systems. We sought to test these possibilities by studying auditory comprehension in acute left versus right hemisphere deactivation during Wada procedures. A series of 20 patients undergoing clinically indicated Wada procedures were asked to listen to an auditorily presented stimulus word, and then point to its matching picture on a card that contained the target picture, a semantic foil, a phonemic foil, and an unrelated foil. This task was performed under three conditions, baseline, during left carotid injection of sodium amytal, and during right carotid injection of sodium amytal. Overall, left hemisphere injection led to a significantly higher error rate than right hemisphere injection. However, consistent with lesion work, the majority (75%) of these errors were semantic in nature. These findings suggest that auditory comprehension deficits are predominantly semantic in nature, even following acute left hemisphere disruption. This, in turn, supports the hypothesis that the right hemisphere is capable of speech sound processing in the intact brain.     

Priming nouns and verbs: differential influences of semantic and grammatical cues in the two cerebral hemispheres

The current experiment investigated how sentential form-class expectancies influenced lexical-semantic priming within each hemisphere. Sentences were presented that led readers to expect a noun or a verb and the sentence-final target word was presented to one visual field/hemisphere for a lexical decision response. Noun and verb targets in the semantically related condition were compared to an unrelated prime condition, which also predicted part of speech but did not contain any lexical-semantic associates of the target word. The semantic priming effect was strongly modulated by form-class expectancy for RVF/LH targets, for both nouns and verbs. In the LVF/RH, semantic priming was obtained in all conditions, regardless of whether the form-class expectancy was violated. However, the nouns that were preceded by a noun-predicting sentence showed an extremely high priming value in the LVF/RH, suggesting that the RH may have some sensitivity to grammatical predictions for nouns. Comparisons of LVF/RH priming to calculations derived from the LSA model of language representation, which does not utilize word order, suggested that the RH might derive message-level meaning primarily from lexical-semantic relatedness.     

The propositional structure of discourse in the two cerebral hemispheres

Readers construct at least two interrelated representations when they comprehend a text: (a) a representation of the explicit ideas in a text and the relations among them (i.e., a propositional representation) and (b) a representation of the context or situation to which a text refers (i.e., a discourse model). In a recent study, found evidence that readers' representations were structured according to propositional relations, but only in the left hemisphere. Both hemispheres, however, appeared to represent contextually relevant semantic information. The goal in the current study was to examine further the organization of explicit text concepts in the two hemispheres. We used an item-priming-in-recognition paradigm in combination with a lateralized visual-field manipulation. We found evidence for a propositionally structured representation in the left hemisphere, that is, priming effects that reflected the linear distance between primes and targets in the propositional structure of passages. We also found that the right hemisphere represented explicit text concepts, but we found no evidence that these concepts were organized structurally. In a second experiment, we found our item priming effects reflected the representation of text information in memory and did not reflect lexical-semantic priming at test.     

Dissociating prelexical and postlexical processing of affective information in the two hemispheres: effects of the stimulus presentation format

Using a lexical decision task, the authors investigated whether brain asymmetries in the detection of emotionally negative semantic associations arise only at a perceptually discriminative stage at which lexical analysis is accurate or can already be found at crude and incomplete levels of perceptual representation at which word-nonword discrimination is based solely on guessing. Emotionally negative and neutral items were presented near perceptual threshold in the left and right visual hemifields. Word-nonword discrimination performance as well as the bias to classify a stimulus as a "word" (whether or not it actually is a word) were assessed for a normal, horizontal stimulus presentation format (Experiment 1) and for an unusual, vertical presentation format (Experiment 2). Results show that while the two hemispheres are equally able to detect affective semantic associations at a prelexical processing stage (both experiments), the right hemisphere is superior at a postlexical, perceptually discriminative stage (Experiment 2). Moreover, the findings suggest that only an unusual, nonoverlearned stimulus presentation format allows adequate assessment of the right hemisphere's lexical-semantic skills.     

Lexical retrieval in left and right brain lesioned children

Performance on two measures of lexical retrieval for 19 left and 13 right brain lesioned children was compared to that of control subjects matched by age, sex, race, and social class. On the Word-Finding Test, left lesioned subjects were significantly slower in response time than left controls when given semantic and visual cues and made more errors when given rhyming cues. On the Rapid Automatized Naming Test, left lesioned subjects were significantly slower than left controls in naming all semantic categories, including colors, numbers, objects, and letters. In contrast, right lesioned subjects responded as quickly as or more quickly than did right controls in all access conditions and in naming semantic categories yet tended to produce more errors than their controls, suggesting a speed-accuracy tradeoff. Children sustaining left brain lesions before 1 year of age appeared to be as impaired as those whose lesions occurred after 1 year of age. Diverse lesion sites within the left hemisphere were associated with increased lexical retrieval latencies.     

Verbal Fluency,Semantics, Context and Symptom Complexes in Schizophrenia

Lexical-semantic access and retrieval was examined in 15 adults diagnosed with schizophrenia and matched controls. This study extends the literature through the inclusion of multiple examinations of lexical-semantic production within the same patient group and through correlating performance on these tasks with various positive and negative clinical symptoms. On tasks of verbal fluency, meaning generation, sentence production using contextual information and confrontation naming, participants with schizophrenia made significantly more semantic errors on naming tasks; produced fewer meanings for homophones; produced fewer items on semantic, phonological, cued and switching fluency tasks; and produced more errors on sentence production tasks when compared to healthy controls. Significant correlations were also observed between ratings of psychomotor poverty and measures of semantic production and mental inflexibility. This study has provided additional evidence for deficits in lexical-semantic retrieval which are not due to underlying semantic store degradation, do not involve phonological based retrieval, and at the level of sentence generation appear to vary as a function of the contextual constraints provided.     

Syntactic, prosodic, and semantic processes in the brain: evidence from event-related neuroimaging

The neural network supporting aspects of syntactic, prosodic, and semantic information processing is specified on the basis of two experiments using functional magnetic resonance imaging (fMRI). In these two studies, the presence/absence of lexical-semantic and syntactic information is systematically varied in spoken language stimuli. Inferior frontal and temporal brain areas in the left and the right hemisphere are identified to support different aspects of auditory language processing. Two additional experiments using event-related brain potentials investigate the possible interaction of syntactic and prosodic information, on the one hand, and syntactic and semantic information, on the other. While the first two information types were shown to interact early during processing, the latter two information types do not. Implications for models of auditory language comprehension are discussed.     

Hemispheric asymmetries in the activation and monitoring of memory errors

Previous research on the lateralization of memory errors suggests that the right hemisphere's tendency to produce more memory errors than the left hemisphere reflects hemispheric differences in semantic activation. However, all prior research that has examined the lateralization of memory errors has used self-paced recognition judgments. Because activation occurs early in memory retrieval, with more time to make a decision, other memory processes, like strategic monitoring processes, may affect memory errors. By manipulating the time subjects were given to make memory decisions, this study separated the influence of automatic memory processes (activation) from strategic memory processes (monitoring) on the production of false memories. The results indicated that when retrieval was fast, the right hemisphere produced more memory errors than the left hemisphere. However, when retrieval was slow, the left hemisphere's error-proneness increased compared to the fast retrieval condition, while the right hemisphere's error-proneness remained the same. These results suggest that the right hemisphere's errors are largely due to activation, while the left hemisphere's errors are influenced by both activation and monitoring.     

The effect of concurrent semantic categorization on delayed serial recall

The influence of semantic processing on the serial ordering of items in short-term memory was explored using a novel dual-task paradigm. Participants engaged in 2 picture-judgment tasks while simultaneously performing delayed serial recall. List material varied in the presence of phonological overlap (Experiments 1 and 2) and in semantic content (concrete words in Experiment 1 and 3; nonwords in Experiments 2 and 3). Picture judgments varied in the extent to which they required accessing visual semantic information (i.e., semantic categorization and line orientation judgments). Results showed that, relative to line-orientation judgments, engaging in semantic categorization judgments increased the proportion of item-ordering errors for concrete lists but did not affect error proportions for nonword lists. Furthermore, although more ordering errors were observed for phonologically similar relative to dissimilar lists, no interactions were observed between the phonological overlap and picture-judgment task manipulations. These results demonstrate that lexical-semantic representations can affect the serial ordering of items in short-term memory. Furthermore, the dual-task paradigm provides a new method for examining when and how semantic representations affect memory performance.     

Hemisphere Inactivation and Mood-State Changes

Mood-state changes were studied during a sodium Amytal testing procedure in epilepsy patients with unilateral temporal lobe foci. The results showed that inactivation of the left hemisphere, but not the right hemisphere, produced a negative mood state. This decline in mood was not specifically related to side of focus or sex. As expected, the inactivation of either hemisphere also lead to an impairment in memory performance. This impairment was somewhat worse during a left than a right hemisphere inactivation. Furthermore, patients with left hemisphere foci showed a lower memory performance than patients with a right-hemisphere lesion. These results suggest (a) a hemispheric specialization for emotion that underlie changes in mood and (b) a discrepancy between mood states at encoding (during inactivation) and retrieval (after inactivation).     

Right hemisphere participation in reading

This study examined whether the right hemisphere's contribution to lexical semantic processing is greatest when it is "disinhibited." Skilled reading may require the controlled modulation of interhemispheric interaction: the left hemisphere (or some other control mechanism) may regulate the subprocesses of reading by selectively "inhibiting" and "disinhibiting" right hemisphere function. Right-handed undergraduates concurrently performed two tasks: a lateralized semantic or rhyme task and a verbal memory task. It was hypothesized that right hemisphere reading processes would be disinhibited when the left hemisphere was "occupied" with the memory task. This hypothesis was supported for a subgroup of subjects who showed evidence of inhibition of right hemisphere function (i.e., left hemisphere dominance for lexical processing) when the lateralized semantic task was performed alone. Across subjects, there was a strong correlation between the degree of left hemisphere dominance in the single-task semantic conditions and the degree of disinhibition of right hemisphere function in the dual-task semantic condition.     

A comparison of lexical-semantic impairments in left hemisphere stroke and Alzheimer's disease

Patients with aphasia due to left hemisphere stroke and patients with Alzheimer's disease, who were matched for severity of naming impairment, were compared on tests of lexical-semantic processing. The results suggest that the lexical-semantic impairments in both groups are due to a combination of impaired access to, and loss of, lexical-semantic information, but that impaired access is more prominent in stroke patients, whereas Alzheimer's disease patients suffer a greater loss of information. The results are discussed in terms of a brain model of the storage and processing of lexical-semantic information, and with respect to implications for treatment strategies.     

Apparent shift in visual field preference after unilateral stroke

Patients with either a left- or a right-hemisphere stroke lesion scored higher in tasks of word-picture matching and of nonverbal shape matching when information was presented tachistoscopically (120 msec) to the visual field (VF) projecting to their undamaged hemisphere. Left-hemisphere stroke patients (n = 13) were dissociated from right-hemisphere stroke patients (n = 15) by low word recognition from memory and by low right VF but nearly normal left VF accuracy in word-picture matching or shape matching; the former appeared to rely upon processing of word meaning by the right hemisphere. In contrast, right-stroke patients had higher right than left VF scores in both tasks, and their discrimination of nonverbal shapes via the right VF was not different from that of controls (n = 15). Preferred processing by the VF projecting to the undamaged hemisphere appeared as a shift in perceptual asymmetry but may indicate, in support of a "direct access" model, that each hemisphere responds more or less efficiently to word and to nonverbal shape discriminations.     

Semantic processing of living and nonliving concepts across the cerebral hemispheres

Studies of patients with category-specific semantic deficits suggest that the right and left cerebral hemispheres may be differently involved in the processing of living and nonliving domains concepts. In this study, we investigate whether there are hemisphere differences in the semantic processing of these domains in healthy volunteers. Based on the neuropsychological findings, we predicted a disadvantage for nonliving compared to living concepts in the right hemisphere. Our prediction was supported, in that semantic decisions to nonliving concepts were significantly slower and more error-prone when presented to the right hemisphere. In contrast there were no hemisphere differences for living concepts. These findings are consistent with either differential representation or processing of concepts across right and left hemispheres. However, we also found a disadvantage for nonliving things compared to living things in the left hemisphere, which is not consistent with a simple representation account. We discuss these findings in terms of qualitatively different semantic processing in right and left hemispheres within the framework of a distributed model of conceptual representation.     

Language acquisition following hemidecortication: Linguistic superiority of the left over the right hemisphere

The language development of three 9- and 10-year-old children possessing only a right or a left hemisphere was studied. Surgical removal of one brain half antedated the beginning of speech, so each child has acquired speech and language with only one hemisphere. Different configurations of language skill have developed in the two isolated hemispheres: phonemic and semantic abilities are similarly developed but syntactic competence has been asymmetrically acquired. In relation to the left, the right brain half is deficient in understanding auditory language, especially when meaning is conveyed by syntactic diversity; detecting and correcting errors of surface syntactic structure; repeating stylistically permuted sentences; producing tag questions which match the grammatical features of a heard statement; determining sentence implication; integrating semantic and syntactic information to replace missing pronouns; and performing judgments of word interrelationships in sentences. Language development in an isolated right hemisphere, even under seizure-free conditions, results in incomplete language acquisition.     

Right Hemisphere Activation during Indirect Semantic Priming: Evidence from Event-Related Potentials

Healthy subjects performed a lexical decision task in a semantic priming paradigm while event-related potentials (ERPs) were recorded from 64 channels. Semantic distance between prime and target was varied by including directly, indirectly, and nonrelated word pairs. At centro-parietal electrodes an N400 to nonrelated pairs was elicited bilaterally which was sensitive only to direct, but not to indirect semantic priming. These N400 priming effects were mirrored by the RT data. At inferior fronto-temporal sites directly related words showed ERP priming effects over both hemispheres. However, indirectly related words only elicited ERP priming effects over the right hemisphere. These results support the hypothesis that the right hemisphere semantic system is involved in processing of remote semantic information.     

Impairments of nonverbal oral movements after left hemisphere damage: A followup analysis of errors

Acquisition of a task requiring the imitation of a series of nonverbal oral movements is most severely impaired in patients with aphasic disturbance. However, nonaphasic patients with left hemisphere damage also perform more poorly than patients with right hemisphere damage or normal control subjects. Analysis of the errors made on the multiple oral movements task reveals that left, but not right, hemisphere damage is highly associated with perseverative responses. Patients with right hemisphere damage do not differ significantly from normal control subjects in the acquisition of or in the type of errors made on the task. It is suggested that the left hemisphere plays an important role in the control of nonverbal oral movement production.