Interhemispheric cooperation and activation in integration of verbal information

Subjects had to judge the semantic relation between two tachistoscopically presented German adjectives. In Exp. I, the two words were projected simultaneously, either both to the left visual field (i.e. to the right hemisphere, LVF-RH), or both to the right visual field (i.e. to the left hemisphere, RVF-LH), or one word to each visual field (i.e. to both hemispheres, EVF-BH). The results yielded a significantly higher performance in the condition EVF-BH than in the two unilateral conditions, without a significant difference between the latter two conditions. In Exp. II, the two words were shown successively. With bilateral presentation, a distinction was made as to whether the left visual field (EVF-BH-L) or the right visual field (EVF-BH-R) was stimulated first. The significantly highest performance was shown in conditions RVF-LH and EVF-BH-R compared with the condition EVF-BH-L, and the significantly poorest performance in the condition LVF-RH compared with the condition EVF-BH-L. The results provide evidence for (1) an intrahemispheric interference with simultaneous unilateral presentation, (2) a cooperative interhemispheric interaction with simultaneous bilateral presentation, and (3) an interhemispheric priming effect with successive bilateral presentation when the left hemisphere was stimulated first.     

Hemispheric semantic priming in the single word presentation task

Functional brain asymmetries in semantic activation were studied by presenting categorically related (e.g. TABLE-BED) or unrelated primes and targets to the left visual field (LVF)/right hemisphere or to the right visual field (RVF)/left hemisphere in the single word presentation lexical decision task. The results showed that the primes in the RVF/left hemisphere primed lexical decisions to the targets both in the RVF and in the LVF. However, the primes in the LVF/right hemisphere did not induce any priming in the LVF or RVF. These results suggest that the left hemisphere automatically activates categorically related meanings in both hemispheres. The role of the right hemisphere in automatic semantic processing may be very limited.     

Task and stimulus demands influence letter-case-specific priming in the right cerebral hemisphere

A greater tendency to complete single-completion word stems (e.g. "BEY") to form previously read whole words (e.g. "BEYOND") was found when test stems were presented in the same letter case as their previously encoded words, compared with the different letter case, but only when stems were presented directly to the right hemisphere (i.e. in the left visual field) and not when they were presented directly to the left hemisphere (i.e. in the right visual field). This finding with single-completion stems was robust (i.e. observed for both lowercase and uppercase stems) when the initial encoding task was perceptually demanding, but it was test-case dependent (i.e. observed for uppercase but not lowercase stems) when the initial encoding task was not perceptually demanding. Results and theory help to explain why letter-case-specific priming in right-hemisphere test presentations is typically test-case dependent when priming is measured using perceptual identification at test, but is consistently robust when priming is measured using multiple-completion word stems (e.g."BEA") at test. Demands from both the stimuli and tasks affect the relative contributions of abstract and specific subsystems to the processing of visual forms.     

Evaluating a split fovea model of visual word recognition: effects of case alternation in the two visual fields and in the left and right halves of words presented at the fovea

Two experiments are reported exploring the effect of cAsE aLtErNaTiOn on lexical decisions to words and nonwords presented laterally or centrally. In line with previous research, Experiment 1 found that case alternation slowed lexical decision responses to words more in the right visual field (RVF) than in the left visual field (LVF). In Experiment 2, the words and nonwords were all presented centrally. There were three conditions, a condition in which the word and nonwords were presented in lower case letters, a condition in which the letters to the left of the central fixation were case alternated (e.g., aMbItion, mOdLants) and a condition in which the letters to the right of fixation were case alternated (e.g., collApSe, pireNtOl). Alternating the case of letters to the right of fixation slowed lexical decision responses more than alternating letter case to the left of fixation. The results provide further support for a split fovea account of visual word recognition according to which those letters of a centrally-fixated word that fall to the left of fixation are processed initially by the right cerebral hemisphere while those letters that fall to the right of fixation are processed initially by the left cerebral hemisphere, with the characteristics of the left and right hemispheres being revealed in the processing of initial and final letters in centrally presented words.     

Opposite hemispheric activations as a result of emotionally threatening and non-threatening words.

Twenty right-handed males participated in a tachistoscopic unilateral letter recognition task with three conditions. In the control condition, each trial consisted of three consonants that were flashed horizontally to the left or to the right visual field. In the threat and non-threat conditions, each lateral three-letter presentation was preceded by an emotionally threatening or non-threatening word presented in central fixation. Across conditions, subjects identified more letters correctly in the right visual field than in the left visual field. The concurrent presentation of threatening words resulted in a selective enhancement of left visual-field performances. The concurrent presentation of non-threatening words resulted in a selective right visual-field enhancement. Our conclusion is that threatening stimuli prime the right hemisphere and can alter predicted laterality patterns.     

Semantic priming by pictures and words in the cerebral hemispheres

In order to determine whether pictures would act as more effective semantic primes than words in the right cerebral hemisphere, automatic semantic activation in intact hemispheres was studied with primed GO-NOGO lexical decision tasks by presenting word-word and picture-word pairs to the left visual field (right hemisphere) or to the right visual field (left hemisphere). Response times in Experiment 1 showed that categorically related targets (e.g., TABLE-BED) were primed only in the right visual field after both word and picture primes. Experiment 2 found that picture primes activated the representations of the corresponding written names in both visual fields. These observations suggest that the range of automatic semantic activation is larger in the left than in the right hemisphere. The results implicate that semantic categories may be organized in a different fashion in the left than the right hemisphere.     

Psychophysics reveals a right hemispheric contribution to body image distortions in women but not men

We tested the hypothesis that the right cerebral hemisphere contributes to the enhanced body image distortions seen in women when compared to men. Using classical psychophysics, 60 right-handed healthy participants (30 women) were briefly presented with size-distorted pictures of themselves, another person (an experimenter), and a non-corporal, familiar object (a coke bottle) to the central, right, and left visual field. Participants had to decide whether the presented stimulus was fatter or thinner than the real body/object, and thus compare the presented picture with the stored representation of the stimulus from memory. From these data we extracted the amount of image distortion at which participants judged the various stimuli to be veridical. We found that right visual field presentations (initial left hemisphere processing) revealed a general "fatter" bias, which was more evident for bodies than for objects. Crucially, a "fatter" bias with own body presentations in the left visual field (initial right hemisphere processing) was only found for women. Our findings suggest that right visual field presentation results in a general size overestimation, and that this overestimation is more pronounced for bodies than for objects. Moreover, the particular "fatter" bias after own body presentations to the left visual field in women supports the notion of a specific role of the right hemisphere in sex-specific body image distortion.     

Cerebral hemispheric differences in memory of emotional and non-emotional words in normal individuals

The present study was designed to examine the cerebral hemispheric differences in memory of positive, negative and non-emotional words using a new method of successive presentation to each visual half-field in which perception of each item was nearly perfect thereby allowing laterality differences for effects of emotion on memory to emerge unconfounded by perception (Experiment 1). This procedure was compared with traditional perceptual identification (Experiment 2). In Experiment 1, 12 words were presented successively in each half field in each trial followed by free recall at the end of the trial. The results showed that recall of positive and negative emotional words was better than that of non-emotional words in both visual fields. Recall of positive and negative emotional words was not different in left and right visual fields (RVFs) although the recall of non-emotional words was better in the RVF than in the left visual field (LVF). The differences in recall between emotional and non-emotional words was greater in the LVF than in the RVF. Experiment 2 used the more traditional method of perceptual identification following each visual half-field presentation of a single item. Perceptual identification was better in the RVF than the LVF in each word condition. There were no visual field differences in perceptual identification between emotional and non-emotional words, as there was for memory in Experiment 1. The results supported the hypothesis that explicit memory for emotional words was dependent more on the right hemisphere, whereas perception of both emotional and non-emotional words was more dependent on the left hemisphere.     

"Bottom-up" and "top-down" effects on reading saccades: a case study

OBJECTIVE: To investigate the role right foveal/parafoveal sparing plays in reading single words, word arrays, and eye movement patterns in a single case with an incongruous hemianopia. METHODS: The patient, a 48-year-old right handed male with a macular sparing hemianopia in his left eye and a macular splitting hemianopia in his right eye, performed various reading tasks. Single word reading speeds were monitored using a "voice-trigger" system. Eye movements were recorded while reading three passages of text, and PET data were gathered while the subject performed a variety of reading tasks in the camera. RESULTS: The patient was faster at reading single words and text with his left eye compared with his right. A small word length effect was present in his right eye but not his left. His eye movement patterns were more orderly when reading text with his left eye, making fewer saccades. The PET data provided evidence of "top-down" processes involved in reading. Binocular single word reading produced activity in the representation of foveal V1 bilaterally; however, text reading with the left eye only was associated with activation in left but not right parafoveal V1, despite there being visual stimuli in both visual fields. CONCLUSIONS: The presence of a word length effect (typically associated with pure alexia) can be caused by a macular splitting hemianopia. Right parafoveal vision is not critically involved in single word identification, but is when planning left to right reading saccades. The influence of top-down attentional processes during text reading can be visualised in parafoveal V1 using PET.     

Emotional words: what's so different from just words?

Aphasic patients, in particular global aphasics, may still swear and produce emotional utterances with ease. Based on these clinical observations we investigated emotional word "reading" in a series of different experiments over 25 years, not only in aphasic patients, but also in the left (LVF) and right (RVF) visual fields of healthy subjects, and in a depth-recorded epileptic patient. Across these experiments we found: i) behaviorally a strong emotional word effect in the left visual field (right hemisphere - RH) of normals, correlating well with the emotional word performance of aphasic patients, pointing to a specific role of the right hemisphere; ii) electro-physiologically a specific early (100-140 msec) brain response to emotional words during scalp recordings in healthy subjects subsequent to right visual field (left hemisphere - LH) stimulation, that source localization procedures project to posterior areas of the right hemisphere; iii) preliminary data of a very early (60 msec) activation of the left amygdala in a depth-recorded epileptic patient when the same emotional words were presented to the right visual field (left hemisphere); and iv) a consistent gender difference showing that the above findings might be relevant for men only. Both hemispheres therefore appear to be implicated in emotional word "reading" but in different ways. We propose that the left amygdala via extrastriate connections acts as a detector of emotional word content at a very early stage of processing; that this amygdala response subsequently modulates the cortical response to emotional words asymmetrically, rendering the left visual cortex less sensitive to emotional words than that of the right hemisphere; and that this modulation is gender dependent.     

Hemispheric asymmetries in memory processes as measured in a false recognition paradigm

Although memory differs in important ways between the left and right cerebral hemispheres, the nature of these differences remains controversial. We examined this issue in two experiments using a false memory paradigm that allowed novel tests of two theories that have not been assessed in a common paradigm previously. Lists of semantically related words (e.g., bed, rest, wake…), all highly associated to one “critical” word (e.g., sleep), were presented auditorily during a study phase. Memory for both the related words and the critical words was measured in a subsequent old/new recognition test using divided-visual- field word presentations. The most important results were that the ability to correctly reject previously unpresented words was greater when test items were presented to the right visual field/left hemisphere (RVF/LH) than to the left visual field/right hemisphere (LVF/RH) and that participants were more confident in correctly rejecting unpresented words when test items were presented to the RVF/LH than to the LVF/RH. Results were in line with the theory that associative activation of semantic information is restricted in the left hemisphere but diffuse in the right; however, these results contrasted with the theory that memory traces are interpretive in the left hemisphere but veridical in the right. A potential resolution to the seemingly contradictory theories of asymmetries in memory processing is briefly discussed.     

Task and Stimulus Demands Influence Letter-case-Specific Priming in the Right Cerebral Hemisphere

A greater tendency to complete single-completion word stems (e.g. “BEY”) to form previously read whole words (e.g. “BEYOND”) was found when test stems were presented in the same letter case as their previously encoded words, compared with the different letter case, but only when stems were presented directly to the right hemisphere (i.e. in the left visual field) and not when they were presented directly to the left hemisphere (i.e. in the right visual field). This finding with single-completion stems was robust (i.e. observed for both lowercase and uppercase stems) when the initial encoding task was perceptually demanding, but it was test-case dependent (i.e. observed for uppercase but not lowercase stems) when the initial encoding task was not perceptually demanding. Results and theory help to explain why letter-case-specific priming in right-hemisphere test presentations is typically test-case dependent when priming is measured using perceptual identification at test, but is consistently robust when priming is measured using multiple-completion word stems (e.g.“BEA”) at test. Demands from both the stimuli and tasks affect the relative contributions of abstract and specific subsystems to the processing of visual forms.     

Hemispheric differences between left and right number representations: effects of conscious and unconscious priming

The contribution of each hemisphere to the generation of number representations was investigated by two lateralized priming experiments in which participants had to compare Arabic digits to a fixed standard of four. In Experiment 1, unmasked primes (Arabic digits or word numerals) were used. In Experiment 2, masked primes were presented consciously or subconsciously. In both experiments similar priming effects were found in the left (LH) and the right hemisphere (RH) when the prime was presented consciously. However, asymmetries emerged when the primes were presented subconsciously: while the priming effects of digits and word numbers were equally large in the right visual half field (RVF-LH), the influence of the word prime on the semantic and the response stage of the left visual half field (LVF-RH) was absent, indicating that a word prime was no longer processed when it was presented subconsciously in LVF-RH. We believe that the origin of the latter effect can be attributed to a failure to transfer word number primes from the RH to the LH when attentional resources are restricted.     

Tapping movements according to regular and irregular visual timing signals investigated with fMRI

Whole-head functional MR images were acquired while 10 subjects were asked to tap with their right index finger in synchrony with a visual stimulus appearing regularly with a frequency of 1.5 Hz, or irregularly with a mean frequency of 1.5 Hz. Performance data show that during regular tapping most taps were close to stimulus onset. However, when the subjects paced their tapping according to the irregular stimuli, most taps appeared about 300 ms after the onset of the pacing stimuli. Comparing the brain activations resulting from regular tapping with those from irregular tapping, we found increased activation in left precuneus only. Comparing irregular versus regular tapping shows increased activity in right cerebellar nuclei and vermis, left ventrolateral thalamus, left sensorimotor cortex, left and right pre-SMA and left SMA proper. These results show that during irregular pacing the motor areas are more strongly activated than during regular pacing. In addition, further neural systems are involved in the motor control during irregular pacing: cerebellar vermis and a cerebello-thalamo-cortical system. The latter is supposedly involved in error correction in the context of visually guided movements.     

Tachistoscopic word recognition performance under conditions of simultaneous bilateral presentation

Four experiments involving the bilateral tachistoscopic presentation of word pairs are reported. This paradigm was found to generate a right hemi-field recognition advantage across manipulations of word orientation, exposure duration, and fixation control. In two of the experiments this recognition asymmetry was maintained in spite of a significant tendency to initiate the verbal report with any available left hemi-field content. In three of the experiments the strength of the left to right order of report tendency was negatively correlated with the magnitude of the right visual field recognition advantage. A hypothesis to account for the more common finding of left hemi-field recognition superiority with bilateral displays is offered.     

Hemispheric asymmetries in feature integration during visual word recognition

Although the definitive source of the left hemisphere's superiority for visual word recognition remains illusive, some argue that the left (LH) and right (RH) hemispheres engage different strategies during early perceptual processes involved in stimulus encoding. In particular, it is proposed that the LH treats a word as a unitary perceptual group whereas the RH processes the letters comprising a word as a series of individual perceptual units. The present study investigated support for this processing distinction by examining hemispheric strategies for temporal integration using Prinzmetal and Millis-Wright's (1984) feature-binding paradigm. A total of 20 participants identified the colour and identity of a target letter, presented within a three-letter word (e.g., ART) or nonword (e.g., HRF), directed to their left or right visual field. Errors were classified on the basis of whether they involved substitution of a colour present within the stimulus but at a different location (ON error), or the substitution of a colour not present within the stimulus (OFF error). As anticipated, for word stimuli there was a higher proportion of OFF errors associated with trials directed to the RH, consistent with the notion that the LH treats words as single perceptual units and is hence biased toward miscombination of perceptual information present within the stimulus. The pattern of ON errors across stimulus type provided clear evidence of RH sequential encoding effects, with the number of errors increasing markedly across the ordinal position of the letters comprising the stimulus string. As such, these data provide new evidence that the LH's advantage for visual word recognition arises, at least in part, from the ability to encode verbal stimuli as single perceptual units.     

Differences in hemispheric asymmetry between dyslexic and normal children on a lateralised lexical decision task

The present experiment was conducted in order to further investigate the relationship between deficits in left hemisphere processing and phonetic decoding in dyslexic children. We administered a lateralised lexical decision task that manipulated wordness, length, and word regularity of grapheme-phoneme conversion. Right-handed male dyslexic children and normal control children were presented with words and pronounceable nonwords. Although there were no overall differences in hemispheric asymmetry between the groups, they did differ in laterality effects in accuracy when responding to nonwords and to phonetically regular words, with the normal children showing the right visual field advantage/ sensitivity (left hemisphere dominance/sensitivity), while the dyslexics failed to show any visual field advantage or sensitivity for these stimuli. Further, group differences were observed in left but not right hemisphere functioning. The results suggest that deficits in left hemisphere processing are apparent only when the dyslexics are attempting to utilise the rules of phonics. Support for the use of this paradigm for use with dyslexic children is also discussed.     

Word repetition within- and across-visual fields: an event-related potential study

A divided visual field (DVF) procedure was used to investigate the scalp distribution of the event-related potential (ERP) repetition effect. ERPs were recorded from 27 scalp sites whilst subjects (n = 20) discriminated between words and non-words presented to either the left (LVF) or the right (RVF) visual field. A proportion of the words were repeated on the trial immediately following their first presentation. In two within-field repetition conditions the two encounters with a word occurred in the same visual field (LVF or RVF). In two across-field repetition conditions, the two encounters with a word occurred in different visual fields. For both words and non-words, task performance was better for RVF presentations than for LVF presentations. In each repetition condition there was a positive-going shift in the ERP elicited by repeated words compared to that elicited by words on their first presentation. This ERP repetition effect was equivalent in magnitude and lateralised to the right hemisphere to an equivalent degree in all four repetition conditions. It is suggested that the ERP effects largely reflect the processing of visual form thought to occur predominately in the right hemisphere.     

Laterality effects for simple and complex geometrical figures, and nonsense patterns

Three groups of normal, right-handed subjects were trained to give same-different responses to pairs of stimuli presented simultaneously to the right or the left side of a fixation point. The subjects exhibited faster reactions (pressing one of two keys) to stimuli appearing in the right visual field when the stimuli were simple geometrical figures (e.g. triangles and squares) or nonsense patterns. By contrast, the subjects showed a left-visual field advantage when the stimuli were complex geometrical figures (e.g. ten- and eleven-sided regular polygons). These differences in performance for the halves of the visual field are attributed to differential hemispheric specializations. The opposite hemispheric superioritieis found with different classes of stimuli are in turn attributed to the discrimination of single features by the left hemisphere and to the use of a spatial strategy by the right hemisphere.     

Interhemispheric cooperation during word processing: evidence for callosal transfer dysfunction in schizophrenic patients

Functional lateralization and interhemispheric interaction during word processing were investigated in schizophrenic patients (n=12) and matched healthy controls (n=18). Words and phonologically regular pseudowords were presented tachistoscopically either in the left or right visual field (unilateral conditions), or simultaneously in both visual hemifields (bilateral condition). Consistent with earlier findings, healthy controls showed a right visual field advantage (RVFA), indicating left-hemispheric dominance for language. The patients showed a RVFA similar to that of controls, consistent with normal left-hemispheric language dominance. Importantly, controls performed much better on words presented in the bilateral condition, when two copies of the same word appeared twice, compared to stimulation in only one of the visual hemifields. This bilateral advantage, which has been interpreted as evidence for cooperation between the hemispheres, was absent in schizophrenics. These data show that schizophrenic patients can exhibit similar lateralization patterns as healthy controls. Their specific functional deficit may be a lack of cooperation between the hemispheres.