The neural basis of the right visual field advantage in reading: an MEG analysis using virtual electrodes

Right-handed participants respond more quickly and more accurately to written words presented in the right visual field (RVF) than in the left visual field (LVF). Previous attempts to identify the neural basis of the RVF advantage have had limited success. Experiment 1 was a behavioral study of lateralized word naming which established that the words later used in Experiment 2 showed a reliable RVF advantage which persisted over multiple repetitions. In Experiment 2, the same words were interleaved with scrambled words and presented in the LVF and RVF to right-handed participants seated in an MEG scanner. Participants read the real words silently and responded "pattern" covertly to the scrambled words. A beamformer analysis created statistical maps of changes in oscillatory power within the brain. Those whole-brain maps revealed activation of the reading network by both LVF and RVF words. Virtual electrode analyses used the same beamforming method to reconstruct the responses to real and scrambled words in three regions of interest in both hemispheres. The middle occipital gyri showed faster and stronger responses to contralateral than to ipsilateral stimuli, with evidence of asymmetric channeling of information into the left hemisphere. The left mid fusiform gyrus at the site of the 'visual word form area' responded more strongly to RVF than to LVF words. Activity in speech-motor cortex was lateralized to the left hemisphere, and stronger to RVF than LVF words, which is interpreted as representing the proximal cause of the RVF advantage for naming written words.     

Hemispheric differences in processing handwritten cursive

Hemispheric asymmetry was examined for native English speakers identifying consonant-vowel-consonant (CVC) non-words presented in standard printed form, in standard handwritten cursive form or in handwritten cursive with the letters separated by small gaps. For all three conditions, fewer errors occurred when stimuli were presented to the right visual field/left hemisphere (RVF/LH) than to the left visual field/right hemisphere (LVF/RH) and qualitative error patterns indicated that the last letter was missed more often than the first letter on LVF/RH trials but not on RVF/LH trials. Despite this overall similarity, the RVF/LH advantage was smaller for both types of cursive stimuli than for printed stimuli. In addition, the difference between first-letter and last-letter errors was smaller for handwritten cursive than for printed text, especially on LVF/RH trials. These results suggest a greater contribution of the right hemisphere to the identification of handwritten cursive, which is likely related visual complexity and to qualitative differences in the processing of cursive versus print.     

Changes in same-different laterality patterns as a function of practice and stimulus quality

Accuracy and reaction time (RT) of judgments about sameness vs. difference of (a) names of two letters and (b) shapes of two nonverbal forms were examined for stimuli presented to the center, left (LVF), and right (RVF) visual fields. For same-name letter pairs during Experiment I, responses were more accurate and faster for LVF than for RVF trials on an initial 90-trial block, but this difference was reversed by a third 90-trial block. The RVF advantage for RT was maintained over Trial Blocks 4 and 5, given during a second session, but had disappeared on Trial Blocks 6 through 9 as RT reached the same asymptotic level for both visual fields. No LVF-RVF differences were obtained at any level of practice for different-name letter pairs or for any of the form pairs. Experiment II replicated the shift from LVF toward RVF advantage that occurred over the first three trial blocks of Experiment I and demonstrated that such a shift does not occur when the letters are perceptually degraded. The results were discussed in terms of differences in cerebral hemisphere specialization for visuospatial vs. abstract stages of letter processing and changes with practice in the relative difficulty of these stages.     

About face: left-hemisphere involvement in processing physiognomies

Three experiments were concluded to investigate the involvement of the two cerebral hemispheres in processing faces. Perceptual discrimination of pairs of faces was equally speedy overall when the stimuli were presented in the right visual field (RVF) or left visual field (LVF). For faces differing in one or two features, however, a qualitatively different pattern of results was obtained for the two visual fields, and an RVF advantage emerged when the difference lay in the upper part of the faces (Experiment 1). An examination of the discriminability of the facial features from which the faces were constructed (Experiment 2) showed that the processes involved in RVF comparisons of faces were not dependent on the saliency of the features but, rather, followed a top-to-bottom serial analysis of the stimuli; the speed of the processing involved in LVF presentations was a function of the degree of similarity of the different comparison faces. Evidence for a serial type of comparison faces were used (Experiment 3). It was concluded that even though comparisons were equally speedy overall in LVF and RVF presentations, qualitatively different processes take place in the two hemispheres, which prove competent at processing faces, each in its own way. Some methodological problems inherent in tachistoscopic studies are discussed, and it is proposed that the quality of the stimulus representation achieved or required for cognitive processing may be determinant in the emergence of functional hemispheric asymmetries.     

Quantitative and Qualitative Hemispheric Asymmetry for Processing Japanese Kana

Native Japanese speakers identified three-letter kana stimuli presented to the left visual field and right hemisphere (LVF/RH), to the right visual field and left hemisphere (RVF/LH), or to both visual fields and hemispheres simultaneously (BILATERAL trials). There were fewer errors on RVF/LH and BILATERAL trials than on LVF/RH trials. Qualitative analysis of error patterns indicated that there were many fewer errors of first-letter identification than of last-letter identification, suggesting top-to-bottom scanning of the kana characters. In contrast to similar studies presenting nonword letter trigrams to native English speakers, qualitative error patterns were identical for the three visual field conditions. Taken together with the results of earlier studies, the results of the present experiment indicate that the ubiquitous RVF/LH advantage reflects a left-hemisphere superiority for phonetic processing that generalizes across specific languages. At the same time, qualitative aspects of hemispheric asymmetry differ from one language to the next and may depend on such things as the way in which individual characters map onto the pronunciation of words and nonwords.     

Is Happy Facial Expression Identified by the Left or Right Hemisphere？

1IntroductionCorrectly identifying other people′s facial ex-pressions of emotions is important to human socialinteraction in all societies.Many studies suggestthat the identification of facial expressions in par-ticular and perceptual processing of emotional infor-mation is carried out mainly by the right hemi-sphere of the brain[1￣7].Damage to the righthemisphere generally produces more significant im-pairment in recognition of all facial expressions ofemotion than damage to the left hemisp…     

Interhemispheric cooperation and non-cooperation during word recognition: evidence for callosal transfer dysfunction in dyslexic adults

Participants report briefly-presented words more accurately when two copies are presented, one in the left visual field (LVF) and another in the right visual field (RVF), than when only a single copy is presented. This effect is known as the 'redundant bilateral advantage' and has been interpreted as evidence for interhemispheric cooperation. We investigated the redundant bilateral advantage in dyslexic adults and matched controls as a means of assessing communication between the hemispheres in dyslexia. Consistent with previous research, normal adult readers in Experiment 1 showed significantly higher accuracy on a word report task when identical word stimuli were presented bilaterally, compared to unilateral RVF or LVF presentation. Dyslexics, however, did not show the bilateral advantage. In Experiment 2, words were presented above fixation, below fixation or in both positions. In this experiment both dyslexics and controls benefited from the redundant presentation. Experiment 3 combined whole words in one visual field with word fragments in the other visual field (the initial and final letters separated by spaces). Controls showed a bilateral advantage but dyslexics did not. In Experiments 1 and 3, the dyslexics showed significantly lower accuracy for LVF trials than controls, but the groups did not differ for RVF trials. The findings suggest that dyslexics have a problem of interhemispheric integration and not a general problem of processing two lexical inputs simultaneously.     

Evaluating a split processing model of visual word recognition: effects of orthographic neighborhood size

The split fovea theory proposes that visual word recognition of centrally presented words is mediated by the splitting of the foveal image, with letters to the left of fixation being projected to the right hemisphere (RH) and letters to the right of fixation being projected to the left hemisphere (LH). Two lexical decision experiments aimed to elucidate word recognition processes under the split fovea theory are described. The first experiment showed that when words were presented centrally, such that the initial letters were in the left visual field (LVF/RH), there were effects of orthographic neighborhood, i.e., there were faster responses to words with high rather than low orthographic neighborhoods for the initial letters ('lead neighbors'). This effect was limited to lead-neighbors but not end-neighbors (orthographic neighbors sharing the same final letters). When the same words were fully presented in the LVF/RH or right visual field (RVF/LH, Experiment 2), there was no effect of orthographic neighborhood size. We argue that the lack of an effect in Experiment 2 was due to exposure to all of the letters of the words, the words being matched for overall orthographic neighborhood count and the sub-parts no longer having a unique effect. We concluded that the orthographic activation found in Experiment 1 occurred because the initial letters of centrally presented words were projected to the RH. The results support the split fovea theory, where the RH has primacy in representing lead neighbors of a written word.     

Lateralized word recognition: assessing the role of hemispheric specialization, modes of lexical access, and perceptual asymmetry

The processing advantage for words in the right visual field (RVF) has often been assigned to parallel orthographic analysis by the left hemisphere and sequential by the right. The authors investigated this notion using the Reicher-Wheeler task to suppress influences of guesswork and an eye-tracker to ensure central fixation. RVF advantages obtained for all serial positions and identical U-shaped serial-position curves obtained for both visual fields (Experiments 1-4). These findings were not influenced by lexical constraint (Experiment 2) and were obtained with masked and nonmasked displays (Experiment 3). Moreover, words and nonwords produced similar serial-position effects in each field, but only RVF stimuli produced a word-nonword effect (Experiment 4). These findings support the notion that left-hemisphere function underlies the RVF advantage but not the notion that each hemisphere uses a different mode of orthographic analysis.     

Error monitoring in the hemispheres: the effect of lateralized feedback on lexical decision.

Does each hemisphere have its own system for monitoring and responding to errors? Three experiments investigate the effect of presenting lateralized accuracy feedback in a bilateral lexical decision task. We presented feedback after each trial in either the left visual field (LVF) or right visual field (RVF). In Experiment 1 the feedback stimuli were faces smiling or frowning, in Experiment 2 we used colored squares, and Experiment 3 tested the effect of verbal feedback. Negative feedback presented in the LVF tended to improve performance on the following trial, while the same negative feedback in the RVF tended to disrupt performance on the following trial. This result was strongest with the faces as feedback, was less pronounced with colored squares, and disappeared with verbal feedback. The results are interpreted as suggesting a right hemisphere superiority for error monitoring that depends on the mode of feedback.     

Hemispheric Differences for Visual Matrix Processing: Stimulus Size and Spatial Frequency Effects

Hemispheric processing differences were assessed by presenting square matrices that varied in size and the number of filled-in cells. Subjects judged whether the matrix contained an even or odd number of filled cells. Experiment 1 employed relatively small matrix sizes (2 x 2, 3 x 3, and 4 x 4), and Experiment 2 employed relatively large matrix sizes (4 x 4, 6 x 6, and 8 x 8). Response time was shorter and error rates lower for left visual field/right hemisphere (LVF/RH) presentations compared to right visual field/left hemisphere (RVF/LH) presentations, with the larger matrices demonstrating the strongest visual field/hemispheric effects. Increases in the number of filled cells contributed to increases for the LVF/RH response time advantage only for the larger arrays. Analysis of the data from both studies collapsed across the number of filled cells produced highly consistent LVF/RH advantages for both response time and error rate, with stronger LVF/RH advantages found for the larger matrix sizes of both studies. The findings suggest that visual stimulus spatial frequency is a key determinant of hemispheric processing advantages, but that this factor is constrained by stimulus size variation. Theoretical implications with respect to the hemispheric processing double filtering by frequency model are discussed.     

Lateralization in the visual perception of Chinese characters and words

It has been reported that tachistoscopic perception of single Chinese characters is better with a left-visual-field (LVF) than with a right-visual-field (RVF) presentation and that of Chinese words consisting of characters is better with a RVF presentation (O. J. L. Tzeng, D. L. Hung, B. Cotton, & S.-Y. Wang, 1979, Nature (London), 382, 499-501). In this study, the nature of this character-word difference in lateralization was explored in a task in which stimuli were presented unilaterally to a visual field for recognition test. Four types of stimuli were used: Single character, single pseudo- or noncharacter, two-character word, and two-character pseudoword. Results show (a) no visual-field advantage for illegal characters and words, (b) a LVF-advantage effect for characters associated with a more prominent LVF than RVF character-superiority effect, (c) a RVF-advantage effect for words associated with a more prominent RVF than LVF word-superiority effect, and (d) these two visual-field effects for characters and words being not absolute, they occur only with a low rather than with a high recognition for their respective illegal counterparts. These results suggest that the character-word difference is due to a more efficient lexical interpretation of character stimuli in the right than in the left hemisphere and a more efficient lexical interpretation of word stimuli in the left than in the right hemisphere.     

Laterality effects in identification of caricatures and photographs of famous faces

Recent evidence suggests that memory representations of familiar faces may exaggerate distinctive information as do caricatures (G. Rhodes, S. Brennan, & S. Carey, Cognitive Psychology, 1987). Therefore caricatures should be effective representations of faces and should yield a right hemisphere processing advantage, as do photographs of faces. Photographs and caricatures of famous faces were presented to the left visual (LVF), the right visual field (RVF), and centrally (CVF), in a name-face verification task. There was a LVF (right hemisphere) advantage for both caricatures and photographs on name-face mismatches but no VF difference for matches. These results were true for both accuracy and reaction time. Processing strategy differences that may account for the difference between matches and mismatches are discussed. Performance was generally better for photographs than for caricatures, irrespective of visual field condition.     

Hemispheric asymmetry and pun comprehension: when cowboys have sore calves

Event-related potentials (ERPs) were recorded as healthy participants listened to puns such as "During branding, cowboys have sore calves." To assess hemispheric differences in pun comprehension, visually presented probes that were either highly related (COW), moderately related (LEG), or unrelated, were presented in either the left or right visual half field (LVF/RVF). The sensitivity of each hemisphere to the different meanings evoked by the pun was assessed by ERP relatedness effects with presentation to the LVF and the RVF. In Experiment 1, the inter-stimulus interval between the pun and the onset of the visual probe was 0 ms; in Experiment 2, this value was 500 ms. In Experiment 1, both highly and moderately related probes elicited similar priming effects with RVF presentation. Relative to their unrelated counterparts, related probes elicited less negative ERPs in the N400 interval (300-600 ms post-onset), and more positive ERPs 600-900 ms post-onset, suggesting both meanings of the pun were equally active in the left hemisphere. LVF presentation yielded similar priming effects (less negative N400 and a larger positivity thereafter) for the highly related probes, but no effects for moderately related probes. In Experiment 2, similar N400 priming effects were observed for highly and moderately related probes presented to both visual fields. Compared to unrelated probes 600-900 ms post-onset, related probes elicited a centro-parietal positivity with RVF presentation, but a fronto-polar positivity with LVF presentation. Results suggest that initially, the different meanings evoked by a pun are both active in the left hemisphere, but only the most highly related meaning is active in the right hemisphere. By 500 ms, both meanings are active in both hemispheres.     

Facilitative orthographic neighborhood effects: the SERIOL model account

A large orthographic neighborhood (N) facilitates lexical decision for central and left visual field/right hemisphere (LVF/RH) presentation, but not for right visual field/left hemisphere (RVF/LH) presentation. Based on the SERIOL model of letter-position encoding, this asymmetric N effect is explained by differential activation patterns at the orthographic level. This analysis implies that it should be possible to negate the LVF/RH N effect and create an RVF/LH N effect by manipulating contrast levels in specific ways. In Experiment 1, these predictions were confirmed. In Experiment 2, we eliminated the N effect for both LVF/RH and central presentation. These results indicate that the letter level is the primary locus of the N effect under lexical decision, and that the hemispheric specificity of the N effect does not reflect differential processing at the lexical level.     

Dividing attention within and between hemispheres: testing a multiple resources approach to limited-capacity information processing

Two experiments tested the limiting case of a multiple resources approach to resource allocation in information processing. In this framework, the left and right hemispheres are assumed to have separate, limited-capacity pools of undifferentiated resources that are not mutually accessible, so that tasks can overlap in their demand for these resources either completely, partially, or not at all. We tested all three degrees of overlap in demand for left hemisphere supplies, using dual-task methodology in which subjects were induced to pay different amounts of attention to each task. Experiment 1 compared complete and partial overlap by combining a verbal memory load with a task in which subjects named nonsense syllables briefly presented to either the left or right visual field (LVF and RVF, respectively). Experiment 2 compared complete versus no overlap by using the same verbal memory load combined with a laterally presented same-different judgment task that did not require a spoken response. Decrements from single-task performance were always more severe when the visual field task stimulus was presented to the RVF. Further, subjects in Experiment 1 were able to trade performance between tasks on both LVF and RVF trials because there was always at least some overlap in left hemisphere demand. In Experiment 2, performance trade-offs were observed on RVF (complete overlap) trials, but not on LVF trials, where no overlap in demand existed. These results contradict a single-capacity model, but they support the idea that the hemispheres' resource supplies are independent and have implications for both cerebral specialization and divided attention issues.     

Strategies of semantic categorization in the cerebral hemispheres

Strategies of semantic categorization in intact cerebral hemispheres were studied in two experiments by presenting names of typical and atypical category instances to the left visual field (LVF) (right hemisphere) or to the right visual field (RVF) (left hemisphere). The results revealed that the typicality of instances had a large effect on categorization times in the LVF in both experiments, suggesting that the right hemisphere relies strongly on a holistic, similarity-based comparison strategy. In Experiment 1, the typicality effect was weaker in the RVF than in the LVF. In Experiment 2, a typicality effect in the RVF was observed for the "four-footed animal" category but not for the "bird" category. The hypothesis that the left hemisphere employs a strategy based on defining or necessary features is not supported by the observed typicality effect in the "four-footed animal" category. Instead, it is suggested that the left hemisphere may be able to categorize on the basis of prestored instance-category knowledge. When such knowledge is not available (e.g., as for four-footed animals), a similarity-based comparison strategy is employed by the left hemisphere.     

Hemispheric differences in the interference among components of compound gratings

The relationship between local/global and high/low spatial-frequency processing in hemispheric asymmetries was explored. Subjects were required to judge the orientation of a high- or low-spatial-frequency component of a compound grating presented in the left visual field (LVF) or right visual field (RVF). In Experiment 1, attention was focused on one or the other component. A signal detection analysis indicated that sensitivity (d′) to the high-spatial-frequency target was reduced more by the presence of the low-spatial-frequency component when both were presented in the LVF rather than in the RVF. In Experiment 2, subjects determined whether a target orientation was present, independent of spatial frequency at only a single level (i.e., at the high- or low-spatial-frequency level), as opposed to both or neither level. An RVF/LH (left hemisphere) advantage was found when the decision was based on the orientation of the high-frequency component. The asymmetrical influence of visual field of presentation and spatial frequency upon sensitivity is discussed in terms of hemispheric differences in the magnitude of inhibition between spatial-frequency channels and in the role of transient channel activity to capture and direct higher order attentional processes.     

Memory scanning of described images and undescribed images: Hemispheric differences

Seamon (1972) has found that reaction times (RTs) to memory probes do not increase with memory set size (M) for words encoded in mental images in a memory scanning task. Rothstein and Atkinson (1975) have failed to replicate Seamon’s results. Experiment 1 investigated this discrepancy by manipulating one methodological difference (whether images were or were not described) between the two investigations. Results revealed that described images produced typical linear increases in RT with M. Undescribed images, however, revealed no change in RT with M for positive probes (i.e., the word presented was contained in the current memory set). Experiment 2 manipulated to which visual field probes were presented (RVF — right visual field; LVF = left visual field). Results showed that the imagery group showed no relation between RT and M for probes presented to the LVF, but a linear relation for probes presentedto the RVF. Described-image and repetition groups revealed a linear relation between RT and M no matter which visual field received the probe.     

Hemisphere dynamics in lexical access: automatic and controlled priming

Hemisphere differences in lexical processing may be due to asymmetry in the organization of lexical information, in procedures used to access the lexicon, or both. Six lateralized lexical decision experiments employed various types of priming to distinguish among these possibilities. In three controlled (high probability) priming experiments, prime words could be used as lexical access clues. Larger priming was obtained for orthographically similar stimuli (BEAK-BEAR) when presented to the left visual field (LVF). Controlled priming based on phonological relatedness (JUICE-MOOSE) was equally effective in either visual field (VF). Semantic similarity (INCH-YARD) produced larger priming for right visual field (RVF) stimuli. These results suggest that the hemispheres may utilize different information to achieve lexical access. Spread of activation through the lexicon was measured in complementary automatic (low probability) priming experiments. Priming was restricted to LVF stimuli for orthographically similar words, while priming for phonologically related stimuli was only obtained in the RVF. Automatic semantic priming was present bilaterally, but was larger in the LVF. These results imply hemisphere differences in lexical organization, with orthographic and semantic relationships available to the right hemisphere, and phonological and semantic relations available to the left hemisphere. Support was obtained for hemisphere asymmetries in both lexical organization and directed lexical processing.