Event-related potential studies of cerebral specialization during reading: I. Studies of normal adults

A number of methodological features were incorporated in a paradigm designed to maximize the likelihood of finding reliable event-related potential (ERP) signs of functional specializations between and within the cerebral hemispheres. Every subject was more accurate in identifying words presented to the right than to the left visual field. The morphology of the ERPs elicited by these words varied considerably as a function of electrode position both within and between the hemispheres. Amplitude asymmetries of ERP components recorded from occipital regions of the two hemispheres varied systematically with the position of the word in the visual field. On the other hand, ERPs from more anterior (temporal and frontal) regions displayed large asymmetries which were in the same direction regardless of the visual field of word presentation. The most prominent such asymmetry was in the negativity in the region 300–500 msec (N410) which was larger in the left than the right hemisphere in every subject. These results demonstrate that in this paradigm which demands specialized language processing ERPs are sensitive to aspects of cerebral organization both within and between the two hemispheres.     

Further evaluation of a sensitive method for detecting experimental brain lesions.

Small, unilateral cortical lesions were placed in either medial-anterior, lateral-anterior, or posterior cortex of 20 male, Sprague-Dawley, albino rats. In addition, unilateral sham operations were performed on 8 rats from the same stock. All subjects had been trained prior to surgery with diametrically opposed visual habits mediated by opposite cerebral hemispheres. The opposing habits were, insofar as possible, evenly matched prior to surgery. After surgery, lesioned hemispheres, regardless of locus of lesion, were slower in acquiring the habit they mediated, than were unlesioned hemispheres. Further, unlesioned hemispheres dominated in test trials in which subjects chose one of the two cue doors with both eyes open. These findings confirm that the "interocularly conflicting discrimination" baseline detects a role of extrastriate cortex in mediation of pattern discrimination habits. They further indicated that losses occur without regard to the various locations of lesion employed.     

Event-related potential studies of cerebral specialization during reading : II. Studies of congenitally deaf adults

A number of methodological features were incorporated in a paradigm designed to maximize the likelihood of finding reliable event-related potential (ERP) signs of functional specializations between and within the cerebral hemispheres. Every subject was more accurate in identifying words presented to the right than to the left visual field. The morphology of the ERPs elicited by these words varied considerably as a function of electrode position both within and between the hemispheres. Amplitude asymmetries of ERP components recorded from occipital regions of the two hemispheres varied systematically with the position of the word in the visual field. On the other hand, ERPs from more anterior (temporal and frontal) regions displayed large asymmetries which were in the same direction regardless of the visual field of word presentation. The most prominent such asymmetry was in the negativity in the region 300–500 msec (N410) which was larger in the left than the right hemisphere in every subject. These results demonstrate that in this paradigm which demands specialized language processing ERPs are sensitive to aspects of cerebral organization both within and between the two hemispheres.     

Interhemispheric visual masking

Backward masking functions were evaluated in a paradigm in which target and masking stimuli were presented to opposite cerebral hemispheres using separate peripheral pathways. Backward masking occurred dichoptically when the target and mask were projected to opposite hemispheres. These findings indicated that some effective masking mechanisms are mediated by interhemispheric interactions independent of retinal phenomena.     

Reversed memory coding and symbol reversal in children

The present study examined an hypothesized mirror-reversed memory-coding phenomenon whereby aligned and mirror-reversed memory records in opposite cerebral hemispheres are confused. Two groups of 7-yr.-old boys were chosen on the basis of tendency to confuse mirror and aligned written symbols. The experimental task required discrimination as same or different of letter pairs which were either same or different and either aligned or mirror-reversed. The pairs were simultaneously presented to right, left, or both cerebral hemispheres. The reversing group had slower reaction times than the normal group in all conditions. Evidence was insufficient to conclude that mirror-same bilateral presentation facilitated responses of either group over aligned-same or peripheral conditions. This result would have been necessary to indicate the presence of a mirror-reversed code across the hemispheres. Research utilizing delayed as well as simultaneous presentation of stimuli to children of various ages would clarify this ambiguity. Bilateral processing across conditions proved easier for both groups, indicating adequate hemispheric integration on a visual-perceptual task even in children who tended to confuse mirror images.     

Hemispheric refractoriness and control of reaction time

Experiments are described which attempt to assess the relative efficiency of the hemispheres and their relationship in performance on complex RT tasks. A divided visual field method was used to direct signals to the temporal or nasal retinae of each eye thus passing information to separate hemispheres. A comparison of the separate response times was used to assess the relative efficiency of each hemisphere but significant differences were not observed. This suggests that each may be the equal of the other in organizing simple responses. A method was used to examine more complex RT by presenting the subject with two simultaneous signals for response. When pairs of signals are directed to separate hemispheres, response times are at their lowest value. When signals are directed to separate hemispheres through the same eye, a significant increase in RT occurs. A source of mutual interference appears to exist at the level of the eye. Response times are extended to their greatest value, however, when both signals are directed to the same hemisphere. This block to function has been described as “hemispheric refractoriness”, and is different for the two hemispheres. While each show a distinct block to function the extent of this is greater in the right or minor hemisphere than it is in the left or major hemisphere.     

Hemisphere differences in conscious and unconscious word reading

Hemisphere differences in word reading were examined using explicit and implicit processing measures. In an inclusion task, which indexes both conscious (explicit) and unconscious (implicit) word reading processes, participants were briefly presented with a word in either the right or the left visual field and were asked to use this word to complete a three-letter word stem. In an exclusion task, which estimates unconscious word reading, participants completed the word stem with any word other than the prime word. Experiment 1 showed that words presented to either visual field were processed in very similar ways in both tasks, with the exception that words in the right visual field (left hemisphere) were more readily accessible for conscious report. Experiment 2 indicated that unconsciously processed words are shared between the hemispheres, as similar results were obtained when either the same or the opposite visual field received the word stem. Experiment 3 demonstrated that this sharing between hemispheres is cortically mediated by testing a split-brain patient. These results suggest that the left hemisphere advantage for word reading holds only for explicit measures; unconscious word reading is much more balanced between the hemispheres.     

Hemispheric Differences in Context Sensitivity During Lexical Ambiguity Resolution

Three experiments were conducted to investigate the influence of contextual constraint on lexical ambiguity resolution in the cerebral hemispheres. A cross-modal priming variant of the divided visual field task was utilized in which subjects heard sentences containing homonyms and made lexical decisions to targets semantically related to dominant and subordinate meanings. Experiment 1 showed priming in both hemispheres of dominant meanings for homonyms embedded in neutral sentence contexts. Experiment 2 showed priming in both hemispheres of dominant and subordinate meanings for homonyms embedded in sentence contexts that biased a central semantic feature of the subordinate meaning. Experiment 3 showed priming of dominant meanings in the left hemisphere (LH), and priming of the subordinate meaning in the right hemisphere (RH) for homonyms embedded in sentences that biased a peripheral semantic feature of the subordinate meaning. These results are consistent with a context-sensitive model of language processing that incorporates differential sensitivity to semantic relationships in the cerebral hemispheres.     

Sulcus topography of the parietal opercular region: an anatomic and MR study

The study describes the sulcal and gyral topography, variability, and left-right asymmetry of the parietal operculum. Eighty postmortem hemispheres as well as sagittal magnetic resonance images from 20 health volunteers (40 hemispheres) were evaluated. Four different types of parietal opercular sulcus topography were recognized. Most frequently, and conforming with the anatomic "textbook pattern", the inferior postcentral sulcus (POCS) is the sulcus anterior to the posterior ascending ramus (PAR) of the Sylvian fissure (type 1). Variations were the following: lack of a PAR (type 2), interposition of an intermediate opercular sulcus and gyrus between PAR and POCS (type 3), and direct transition of PAR into POCS with subsequent lack of a classical supramarginal gyrus (type 4). Inconstancy of the sulcal standard arrangement was especially pronounced among left hemispheres, where the patterns differed from type 1 in one third of the cases. Types 2 and 3 were significantly more frequent in left hemispheres, whereas type 4 occurred significantly more frequently in right hemispheres. Upon intraindividual left-right comparison, a remarkable 38% of the brains showed gross asymmetry of the parietal opercular sulcus patterns, characterized by a left type 2 or 3 and/or a right type 4; another 5% exhibited a reverse type of asymmetry. The findings supplement previous data on gross variability and left-right asymmetry of the posterior Sylvian fissure and its lower bank. They indicate that the Sylvian fissure is an unreliable landmark with respect to inferior parietal structures especially in left hemispheres. Individual mapping of perisylvian topography may contribute to studies on structural-functional relationship.     

Facilitation and disruption of lateralized syllable processing by unattended stimuli in the opposite visual field

The influence of lateralized unattended stimuli on the processing of attended stimuli in the opposite visual field can shed light on the nature of information that is transferred between hemispheres. On a cued bilateral task, participants tried to identify a syllable in the attended visual field, which elicits a left hemisphere (LH) advantage and different processing strategies by the two hemispheres. The same or a different syllable or a neutral stimulus appeared in the unattended field. Transmission of unattended syllable codes between hemispheres is symmetric, as revealed by equal interference for the two visual fields. The LH is more accurate than the RH in encoding unattended syllables, as indicated by facilitation in the left but not right visual field and a greater frequency of identifiable intrusions into the left than right field. However, asymmetric encoding strategies are different for attended and unattended syllables.     

Hemispheric differences in picture-word interference

In a picture-word version of the Stroop task, 30 right-handed subjects were tested under each of six conditions in which a picture alone or a picture plus a word were presented to the left, the right, or both hemispheres. In two additional conditions the picture was presented to the right hemisphere and the word was simultaneously presented to the left hemisphere, or vice versa. For all conditions, subjects were instructed to name the picture only, as rapidly as possible. Picture naming times were significantly slower for the conditions in which the pictures were accompanied by words than in the respective picture alone conditions. Moreover, simultaneous presentation of a picture and a word to both hemispheres resulted in greater interference (slower picture naming times) than did the simultaneous presentation of the picture and the word to either the left hemisphere alone or the right hemisphere alone. The latter two conditions, in turn, resulted in significantly more interference than did the simultaneous presentation of the picture to one hemisphere and the word to the other hemisphere. This pattern of results suggests that the Stroop effect obtained under normal circumstances is in large part a function of the interference caused by the simultaneous processing of items in the same hemisphere. In contrast to hemispheric differences reported for the color-word Stroop task, the effect of presenting a picture and word simultaneously to the right hemisphere did not differ reliably from that of presenting a picture and word to the left hemisphere. The failure to replicate this aspect of the color-word Stroop is attributed to differences in the abilities of the two hemispheres to process the respective target items (the color or the picture) of the two tasks.     

Aging and Individual Variation in Interhemispheric Collaboration and Hemispheric Asymmetry

ABSTRACT

Thirty younger (Mean Age = 19.9 years) and 20 older adults (Mean Age = 74.7 years) performed Physical and Name Identity letter-matching tasks (matches were either within or between hemispheres) to study age-related changes in 1) the efficiency with which the two hemispheres interact with each other and 2) hemispheric asymmetry. In order to determine whether age-related effects were associated with differences in cognitive resources, the same individuals completed a set of memory span tasks. Performance on the letter-matching tasks indicated that the costs of interhemispheric collaboration were greater for older than for younger participants. However, within the older group, the advantage of spreading processing across both hemispheres increased as memory span decreased, suggesting that older individuals who are challenged by cognitive complexity are more likely to show increased benefits from between-hemisphere processing than individuals who are not so challenged. There was also an overall left visual field/right hemisphere advantage for the younger but not for the older group, suggesting greater age-related declines in right- than left-hemisphere function.     

Hemispheric specialization for temporal information: Implications for the perception of voicing cues during speech perception

Developmental research reporting electrophysiological correlates of voice onset time (VOT) during speech perception is reviewed. By two months of age a right hemisphere mechanism appears which differentiates voiced from voiceless stop consonants. This mechanism was found at 4 years of age and again with adults.A new study is described which represents an attempt to determine a more specific basis for VOT perception. Auditory evoked responses (AER) were recorded over the left and right hemispheres while 16 adults attended to repetitive series of two-tone stimuli. Portions of the AERs were found to vary systematically over the two hemispheres in a manner similar to that previously reported for VOT stimuli. These findings are discussed in terms of a temporal detection mechanism which is involved in speech perception.     

Intrahemispheric differentiation of vowels: Principal component analysis of auditory evoked responses to computer-synthesized vowel sounds

Auditory evoked brain responses (AER) were recorded in response to a series of synthesized vowel sounds which varied in formant bandwidth. Multivariate analyses indicated that changes in AER component structure recorded from different scalp regions over both hemispheres varied as a function of different vowel sounds and formant bandwidth. No interhemispheric differences in scalp AER distributions were noted.     

Role of input factors in visual-field asymmetries

This paper examines the implications of lateral tachistoscopic presentation of information for the processing efficiency of the intact cerebral hemispheres. Considering that the understanding of the processes underlying the particular competences of each hemisphere may require, as a preliminary step, the specification of the characteristics of the input on which the brain operates, anatomical, physiological, and psychophysical consequences of briefly stimulating the retinal periphery for the representation of information in the brain are outlined, with special reference to the spatial-frequency spectral composition of the stimuli. Retinal eccentricity and brief exposure duration converge to making the representation of information qualitatively different from the information that the brain normally operates on, which constrains the interpretation of findings with respect to the normal functions of the cerebral hemispheres. A review and discussion of empirical findings relevant to these issues suggest that manipulation of procedural variables may differentially affect the processing efficiency of the cerebral hemispheres and indicate that a given pattern of visual-field asymmetry may be overdetermined by a multitude of variables interacting in complex ways.     

Hemispheric alpha asymmetries of stuttering males and nonstuttering males and females for words of high and low imagery

Alpha hemispheric asymmetries of nonstuttering males, nonstuttering females, and stuttering males were explored with electroencephalographic procedures during exposure to two lists of one-syllable words which the subjects were required to recall following presentation. One word list contained low-imagery words while the other contained high-imagery words. Stuttering males were found to have significantly less alpha in their right hemispheres, suggesting right hemispheric processing strategies. This finding was interpreted as supporting the “segmentation dysfunction” explanation of stuttering suggested by Moore and Haynes (1979b). Nonstuttering males showed less left than right alpha while nonstuttering females revealed no difference between their right and left hemispheres. Differences between nonstuttering males and females are discussed as being task and stimuli dependent relative to the segmental/nonsegmental nature of both task and stimuli. Differential hemispheric asymmetries for words of high or low-imagery were not observed.     

Interhemispheric categorization of pictures and words

Earlier studies suggest that interhemispheric processing increases the processing power of the brain in cognitively complex tasks as it allows the brain to divide the processing load between the hemispheres. We report two experiments suggesting that this finding does not generalize to word-picture pairs: they are processed at least as efficiently when processed by a single hemisphere as compared to processing occurring between the two hemispheres. We examined whether dividing the stimuli between the visual fields/hemispheres would be more advantageous than unilateral stimulus displays in the semantic categorization of simultaneously presented pictures, words, and word-picture pairs. The results revealed that within-domain stimuli (semantically related picture pairs or word pairs) were categorized faster in bilateral than in unilateral displays, whereas cross-domain stimuli (word-picture pairs) were not categorized faster in bilateral than in unilateral displays. It is suggested that interhemispheric sharing of word-picture stimuli is not advantageous as compared to unilateral processing conditions because words and pictures use different access routes, and therefore, it may be possible to process in parallel simultaneously displayed word-picture stimuli within a single hemisphere.     

汉字属性匹配与大脑两半球协同活动

在识别汉字时,大脑两半球同时加工的作业成绩是否随加工阶段的增加而显出优势。本研究用汉字形、音、义三种属性分别相匹配的方法用用指定指定提取法,所得实验结果一致表明,形似材料双侧视野同时识别与单侧视野识别成绩无显著差别,而同音与近义材料则表现出双侧呈现显著优于单侧呈现的作业成绩。最后结合脑功能成像研究,对加工阶段影脑半球佃同活动的原因进行了讨论。     

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.     

Visual and verbal coding in the interhemispheric transfer of information

A manual response was used to record a decision of ‘same’ or ‘differ’ on two visual signals which were presented simultaneously, either both to the same hemisphere or each to different hemispheres. The signals were drawn from the set of letters A, B, a, b. Eight subjects were required to match on the basis of the names of the letters (irrespective of whether they were capitals or lower case) and eight subjects were required to match on the basis of the visual identity of the signals.Reaction times were measured for ‘same’ and ‘differ’ decisions.This experiment extends a previous study by the same authors (Davis and Schmit 1971). The previous finding, that when two signals are presented each to different hemispheres, reaction times are shorter than when both are presented to the same hemisphere, was confirmed for both visual and name matching. New evidence is provided on the role of each hemisphere in the analysis and comparison of signals, on either a visual or a verbal basis, for ‘same’ and ‘differ’ decisions. A model to explain the results in terms of the different functions of the two hemispheres is proposed.