Right hemisphere sensitivity to arousal and depression.

Several lines of evidence show impaired right hemisphere function in depression. Lateralized simple reaction time tasks show impaired left visual field responses both in normals experiencing a depressed mood and in patients with mild unipolar depression. One interpretation for these findings is that depression impairs right hemisphere function by interfering with right hemisphere arousal and vigilance mechanisms. In order to test this hypothesis, subjects receiving either depression or relaxation mood suggestions performed an uncued reaction time task that has been shown to be sensitive to right posterior brain damage. Level of alertness was varied by contrasting uncued blocks with blocks in which targets were preceded by a warning tone. The results showed the predicted slowing of left visual field responses in the depressed mood, but only in women. The effect was significant only for the uncued blocks. The left visual field impairment was significantly larger during depression than in the relaxation state, but a smaller left visual field slowing was present in women in the relaxed state as well. These results may be consistent with the notion that depression interferes with right hemisphere function in part by influencing right hemisphere arousal mechanisms.     

Hemisphere differences in the mood state-dependent effect for recognition of emotional faces

Forty subjects viewed 10 pictures of facial expressions of emotion while they experienced a happy mood and 10 pictures while they experienced a sad mood. Later, while re-experiencing either a happy or sad mood, they were tested for recognition of these 20 target pictures intermixed with 20 distractors. Recognition of the 10 pictures seen earlier in a disparate mood was impaired significantly when they were presented at testing to the right hemisphere, but not when presented to the left. The right hemisphere appears to store the subject's mood as an integral part of a memory representation for an emotionally expressive face. When that face is presented at testing to the right hemisphere, recognition depends on whether the subject's test mood matches the mood stored in the representation. In contrast, the left hemisphere appears to store the subject's mood separately from encoded visual information about a face, and so recognition of a face presented at testing to the left hemisphere is unaffected by changes in mood.     

Hemispheric specialization for emotional expression: a reexamination of results from intracarotid administration of sodium amobarbital

Emotional and behavioral responses to sodium amobarbital injection were examined in 44 epilepsy surgery candidates. Thirty-three emotional reactions occurred in 26 patients, 44% of the reactions following right hemisphere injection and 32% after injection of the left hemisphere. Among all patients examined, laughter/elated mood was statistically more frequent following right hemisphere injection while crying was statistically associated with left hemisphere injections. These observed effects were not related to amobarbital dose, sex, bilateral hemispheric inactivation (angiographic crossflow), general cognitive status, or side of seizure onset. These results support differing specialization of emotional expression in the right and left cerebral hemispheres and their subcortical connections.     

Hemispheric processing asymmetries: implications for memory

Recent research has demonstrated that memory for words elicits left hemisphere activation, faces right hemisphere activation, and nameable objects bilateral activation. This pattern of results was attributed to dual coding of information, with the left hemisphere employing a verbal code and the right a nonverbal code. Nameable objects can be encoded either verbally or nonverbally and this accounts for their bilateral activation. We investigated this hypothesis in a callosotomy patient. Consistent with dual coding, the left hemisphere was superior to the right in memory for words, whereas the right was superior for faces. Contrary to prediction, performance on nameable pictures was not equivalent in the two hemispheres, but rather resulted in a right hemisphere superiority. In addition, memory for pictures was significantly better than for either words or faces. These findings suggest that the dual code hypothesis is an oversimplification of the processing capabilities of the two hemispheres.     

Left hemispheric interference with nonverbal performance in aphasics: comparison with data from split-brain studies

Two nonverbal tasks (Nebes' Figural Unification Test and Arc-Circle Matching Task) were given to 10 aphasic patients and 10 normal controls. Observations in split-brain patients have shown that the right hemisphere performs almost normally on these tasks while the left hemisphere fails completely. Left and right hand performances of aphasics and controls were compared with those of split-brain patients as reported in the literature. In the Figural Unification Test the aphasics' left hand performance was significantly poorer than that of split-brain patients. This is interpreted as an effect of left hemispheric interference with right hemispheric nonverbal performance. The lack of a similar impairment of left hand performance in the Arc-Circle Matching Task suggests that this interference is only provoked by tasks which, due to the possible verbalization of stimulus material, tend to stimulate left hemispheric activity.     

Processing persuasive arguments: recall and recognition as a function of agreement and manipulated activation asymmetry

Previous research has demonstrated greater attention toward disagree arguments and a greater persuasibility during relative right than during relative left hemisphere activation. This experiment was performed to conceptually replicate and extend these findings to memory stages in the processing of persuasive arguments. The experiment used induction of lateral orientation of attention as the means to selectively activate regions of the brain in normal right-handed subjects. The results showed the expected significant interaction for correct recall as a function of agreement with the argument and hemisphere activated. There was greater recall of agree arguments during left hemisphere activation and greater recall of disagree arguments during right hemisphere activation, as predicted. The effect for recognition memory was in the same direction, but was statistically nonsignificant. These findings support previous research on cerebral asymmetries in the processing of persuasive arguments. This phenomenon is discussed as a possible outcome of asymmetries of affect.     

Regional cerebral blood flow response to recognition memory versus semantic classification tasks

Two randomly assigned groups, N = 7 each, of neurologically normal patients were composed. One group, RM, had regional cerebral blood flow measurements during a recognition memory probe test; the other group, SC, had rCBF measurements during a semantic classification task, using the same stimulus and response features as the RM task. Thus, the groups differed only in the cognitive instruction they were executing. The RM group showed a significantly lower change from resting baseline to activation, in mean left hemisphere initial slope index, than did the SC group. The RM group, but not the SC group, showed a significant inverse correlation between occipital flow and accuracy of memory performance as indicated by d′. Correlations between age and hemispheric initial slope index, and between homologous left and right hemisphere sites, were also described. The results are considered to support an anatomical basis for the distinction between episodic and semantic memory and to suggest that occipital flow may diminish with accurate memory performance because of an upstream demand of blood in the medial temporal lobes.     

Role of amygdala-prefrontal cortex circuitry in regulating the expression of contextual fear memory

The basolateral amygdala (BLA) and medial prefrontal cortex (mPFC) are inter-connected regions involved in fear memory expression. The reciprocal nature of projections between these areas differs along the rostrocaudal extent of BLA. This study investigated the role of functional interactions between BLA and the prelimbic (PL) subregion of mPFC in mediating contextual fear memory. Freezing served as the measure of conditioned fear. Experiments 1–3 examined the effects of left, right or bilateral infusion of bupivacaine into anterior BLA (aBLA), posterior BLA (pBLA) or PL on fear memory expression. Reversible inactivation of left, right or bilateral aBLA impaired fear memory expression. Bilateral inactivation of pBLA or PL also disrupted the expression of fear memory, although left or right inactivation alone had no significant effects in either region. Experiment 4 examined the effects of functionally disconnecting pBLA and PL on contextual fear memory by infusing bupivacaine unilaterally into pBLA and PL in the ipsilateral or contralateral hemisphere. Fear memory expression was impaired by asymmetric inactivation of pBLA and PL; however, a similar effect was also observed with symmetric inactivation of these regions. Bupivacaine infusion did not affect behavior in the open field, likely ruling out non-specific effects of inactivation on innate fear and locomotor activity. These results demonstrate different roles for rostral and caudal BLA in mediating the expression of contextual fear memory. They also raise the possibility that pBLA–PL circuitry is involved in subserving fear memory expression via complex processing mechanisms, although further research is needed to confirm this preliminary finding.     

Controlled recall of verbal material in temporal lobe epilepsy

This study used a guided process‐dissociation procedure to examine the contribution of controlled and automatic uses of memory to a cued‐recall task in 24 patients with unilateral temporal lobe epilepsy (TLE: 12 left‐sided; 12 right‐sided), and 12 neurotypical controls. In an inclusion task, participants attempted to complete three‐letter word stems using previously studied words, in an exclusion task they aimed to avoid using studied words to complete stems. Patients with left TLE produced fewer target completions under inclusion conditions. Completion rates were not significantly different under exclusion conditions. Estimates derived from process dissociation calculations, confirmed that the cued‐recall deficit in left TLE patients arose entirely from impairment in controlled memory processes. There were no group differences in the estimates of automatic processes. Recognition judgements of stems corresponding to studied words did not differ between the groups. Overall the results support the view that controlled and automatic memory processes are mediated by separable neural systems. Hippocampal and related structures within the left MTL are more important than corresponding right hemisphere structures for the controlled retrieval of verbal material. In contrast, the findings from this study do not suggest that the left and right temporal lobes make a differential contribution to automatic memory processing. The theoretical and clinical relevance of these findings are discussed.     

Testing conclusions from functional imaging of working memory with data from acute stroke

Functional imaging studies indicate that the left hemisphere mediates verbal working memory, while the right hemisphere mediates both verbal and spatial working memory. We evaluated acute stroke patients with working memory tests and imaging to identify whether unilateral dysfunction causes deficits in spatial and/or verbal working memory deficits. While left cortical stroke patients had verbal working memory impairments (p<0.003), right cortical stroke patients had both verbal p<0.007) and spatial working memory (p<0.03) impairments, confirming functional imaging results. Patients with transient ischemic stroke and patients with non-cortical stroke did not have significant deficits in working memory in either modality.     

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.     

Imagery and verbal encoding in left and right hemisphere damaged patients

Neuropsychological aspects of imaginal and verbal encoding in memory were explored in two forced-choice recognition memory experiments with patients suffering from left and right anterior cerebral hemisphere damage. In the first experiment stimulus type and rate of presentation were varied. Predictions of patient performance based on the hypothesis that left anterior hemisphere pathology impairs verbal memory coding and right anterior hemisphere pathology impairs imaginal coding were confirmed. In a second recognition memory experiment for pictures of common objects, system-specific (imaginal or verbal) interference and distractor effects were demonstrated by analyzing the effects of interpolated tasks and the nature of false-recognition errors.     

Laterality of lesions and trait-anxiety on working memory performance

An asymmetry of anterior cerebral activation favoring the right hemisphere has been associated with dispositional negative affect including trait-anxiety, while the opposite appears true of cerebral asymmetry favoring the left hemisphere. It was hypothesized that an asymmetry of cerebral activation, as defined by scores on a measure of trait-anxiety, ipsilateral to the side of an anterior brain lesion would be associated with less efficient cognitive processing than greater activation in the hemisphere contralateral to the lesion. Patients with anterior left (n = 16) or right (n = 15) hemisphere lesions completed the State-Trait Anxiety Inventory and several neurocognitive tasks. Of the abilities tested, only Digit Span scores showed an interaction between side of lesion and presumed activation asymmetry. Patients with right- but not with left-hemisphere damage showed significant differences in working memory performance depending on the presumed direction of asymmetry of the two hemispheres, supporting the dual roles of the right hemisphere in affective processing and directed attention.     

Form-Specific Visual Priming in the Left and Right Hemispheres

Word fragment completion performance was examined for items that were presented in the same or different letter case at study and test. During the study phase words and nonwords were presented at central fixation, then during the test phase a divided visual field technique was used in which word fragments were presented briefly to the right hemisphere (left visual field) or the left hemisphere (right visual field). Previous research using the word stem completion task indicated that only the right hemisphere was sensitive to case changes in words from study to test. In contrast, the current results indicate that in the fragment completion task the priming effects for the test items presented to either hemisphere were greater when the fragments were in the same compared to different letter case at study and test. These results indicate that both hemispheres are capable of supporting form-specific visual implicit memory.     

Unilateral brain lesions in childhood: performance on the Revised Token Test

Comprehension of connected language as assessed by the Revised Token Test (RTT) was studied in 17 children with left hemisphere lesions (LL) and 11 with right hemisphere lesions (RL). LL children's significantly lower performance on several subtests than left controls (LC) matched by age, sex, race, and social class appeared to be related to the memory demands of these subtests rather than the limited syntactic elements assessed by the RTT. LL subjects requested significantly more command repetitions than control or RL subjects and when a repetition was not requested LL subjects often responded with immediacy. Although RL children tended to perform lower than right matched controls (RC), these differences were not significant nor readily related to either the memory or specific linguistic structures assessed. RL subjects required fewer command repetitions than LL subjects and exhibited a significantly greater frequency of immediate responses than controls or LL subjects. While based on few children, a trend for LL children with retrorolandic lesions to perform more poorly than those with left prerolandic lesions was suggested. No systematic difference in performance was apparent for children with left cortical vs. left subcortical lesions or among discrete sites of lesions within the right hemisphere. Children with left lesions prior to 1 year of age performed no better and, in several instances, significantly poorer than LL patients sustaining lesions after 1 year of age. Among RL subjects, those sustaining lesions after 1 year of age had greater difficulty than those with lesion onset before 1 year of age, especially on linguistic elements which seemed to be dependent upon visual spatial properties. Further studies are needed to evaluate the comprehension of more complex linguistic structures among children with unilateral brain lesions as well as to study the role of more basic factors such as memory and attention in explaining the present findings.     

Hemispheric processing of inferences: The effects of textual constraint and working memory capacity

In this study, we investigated hemispheric differences in the generation of bridging and predictive inferences. Participants read texts that provided either strong or weak causal constraints for a particular bridging (Experiment 1) or predictive (Experiment 2) inference and performed a lexical decision task to inference-related targets presented to the left or the right hemisphere. Facilitation for strongly constrained bridging and predictive inferences was found in both hemispheres. In contrast, facilitation for weakly constrained inferences was stronger in the right than in the left hemisphere for both bridging and predictive inferences, although for the latter there was some facilitation in the left hemisphere as well. We also considered whether these effects differ as a function of the working memory capacity of the reader. High working memory capacity readers showed greater facilitation for strongly constrained inferences than for weakly constrained inferences in both hemispheres, whereas low working memory capacity readers showed this same pattern in the left hemisphere but showed equal facilitation for strongly and weakly constrained inferences in the right hemisphere. These results suggest that hemispheric processing, textual constraint, and working memory capacity interact to affect how readers generate causal inferences.     

Duration and numerical estimation in right brain‐damaged patients with and without neglect: Lack of support for a mental time line

Previous studies have shown that left neglect patients are impaired when they have to orient their attention leftward relative to a standard in numerical comparison tasks. This finding has been accounted for by the idea that numerical magnitudes are represented along a spatial continuum oriented from left to right with small magnitudes on the left and large magnitudes on the right. Similarly, it has been proposed that duration could be represented along a mental time line that shares the properties of the number continuum. By comparing directly duration and numerosity processing, this study investigates whether or not the performance of neglect patients supports the hypothesis of a mental time line. Twenty‐two right brain‐damaged patients (11 with and 11 without left neglect), as well as 11 age‐matched healthy controls, had to judge whether a single dot presented visually lasted shorter or longer than 500 ms and whether a sequence of flashed dots was smaller or larger than 5. Digit spans were also assessed to measure verbal working memory capacities. In duration comparison, no spatial‐duration bias was found in neglect patients. Moreover, a significant correlation between verbal working memory and duration performance was observed in right brain‐damaged patients, irrespective of the presence or absence of neglect. In numerical comparison, only neglect patients showed an enhanced distance effect for numerical magnitude smaller than the standard. These results do not support the hypothesis of the existence of a mental continuum oriented from left to right for duration. We discuss an alternative account to explain the duration impairment observed in right brain‐damaged patients.     

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.     

The impact of left and right intracranial tumors on picture and word recognition memory

This study investigated the effects of left and right intracranial tumors on picture and word recognition memory. We hypothesized that left hemispheric (LH) patients would exhibit greater word recognition memory impairment than right hemispheric (RH) patients, with no significant hemispheric group picture recognition memory differences. The LH patient group obtained a significantly slower mean picture recognition reaction time than the RH group. The LH group had a higher proportion of tumors extending into the temporal lobes, possibly accounting for their greater pictorial processing impairments. Dual coding and enhanced visual imagery may have contributed to the patient groups' similar performance on the remainder of the measures.