Hemispheric asymmetries in human infants: Spectral analysis of flash and click evoked potentials

Evoked potentials of 16 human infants (mean age = 5.0 weeks, SD = 1.8 weeks) were recorded from the left and right, occipital and temporal areas. Spectral analysis showed a high amplitude, localized, coherent center of activity within the left temporal area for click stimuli, and a high amplitude, localized center of activity in the right occipital area for flash stimuli. It was proposed that the structured auditory information of the click and the unstructured visual information of the flash represented different degrees of familiarity to the subjects. With this hypothesis, left hemisphere involvement in stimulus processing would increase as the stimulus became more referrable to previous long- or short-term experience. Conversely, right hemisphere involvement would increase with unfamiliar stimuli which could not be readily associated with earlier data.     

Right-hemisphere responses from preschool children to temporal cues to speech and nonspeech materials: electrophysiological correlates

Auditory-evoked responses (AERs) were recorded from scalp electrodes placed over the left and right temporal hemisphere regions of 12 preschool children while they listened to a series of velar stop consonants which varied in voice onset time (VOT) and to two-formant tone stimuli with temporal lags comparable to the speech materials. A late occurring negative peak (N400) in the right hemisphere AERs discriminated between both the speech and nonspeech materials in a categorical-like manner. Sex-related hemisphere differences were also noted in response to the two different stimulus types. These results replicate earlier work with speech materials and suggest that temporal delays for both speech and nonspeech auditory materials are processed in the right hemisphere.     

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

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

The processing of fundamental frequency in a dichotic matching task

The temporal characteristics of auditory memory were investigated using a reaction time paradigm. The study attempted to determine whether or not there are functionally distinct left and right hemisphere components of auditory memory with different capacities for the retention of nonlinguistic information, such as fundamental frequency, over the course of several seconds. The results indicated that fundamental frequency information in auditory memory remains substantially unchanged over the course of 2 sec. No evidence was found for the existence of separate left and right hemisphere components of auditory memory with different temporal characteristics. The implications of this finding for the organization of auditory memory in the brain are discussed.     

Deficits in temporal sequencing of verbal material: The effect of laterality of lesion

Temporal sequencing of verbal materials (digits, words, and geometric forms) presented in two sensory modalities (auditory and visual) to three groups of subjects (Broca's with left anterior lesions, patients with right hemisphere lesions, and normals) was examined. Each subject was asked to point to a set of stimuli in the same sequence as presented by the examiner. Results indicated that patients with left hemisphere lesions were more impaired on all tasks than the right hemisphere lesioned patients who, in turn, were impaired compared to normal controls. Response to auditory presentation was superior to response to visual presentation. Also, digits were the easiest for all groups, and words were easier than geometric forms. Of special interest was the finding which suggested that right hemisphere lesions are associated with impairment of verbal temporal sequencing under either auditory or visual presentation.     

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

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

Cerebral control for speech in right-handers and left-handers: an analysis of the views of Paul Broca, his contemporaries, and his successors

According to several recent historical accounts, Broca (1865a) stated that left-handers are the mirror-reverse of right-handers for cerebral control of speech, with the right hemisphere being dominant in left-handers, and the left hemisphere dominant in right-handers. The same accounts then note Broca's error in light of current evidence that the majority of left-handers are left-dominant for speech just as are nearly all right-handers. Eling (1984) has called such statements misrepresentations of Broca's position and has argued that Broca's analysis actually was more compatible with the current view that there is a disjunction, meaning an absence of an intimate anatomical relationship, between cerebral control for handedness and speech. The current paper looks again at Broca's work, describes the context in which his views were first articulated, and traces the development of the mirror-reversal principle. The conclusion is reached that, judged by a narrow reading of the 1865 paper, Broca's views could indeed be construed as an anticipation of the modern disjunction principle. However, judged by a broader reading, by consideration of his other writing, and in the context of the philosophical and scientific tradition that shaped his work, it is suggested that it was the mirror-reversal principle to which Broca was actually disposed.     

Persisting apraxia in two left-handed, aphasic patients with right-hemisphere lesions

Apraxia usually follows a left hemisphere lesion in right-handers with left hemisphere speech representation. Apraxia following a right hemisphere lesion in left-handers is rare, however, and not well documented in the literature. Two left-handed patients are described in whom apraxia and aphasia followed a right hemisphere lesion. Both the apraxic and the aphasic deficits improved but were still demonstrable 6 weeks following the infarct. The data are consistent with those for right-handers with left hemisphere lesions in suggesting some overlap of anatomical structures for the control of speech and praxis.     

Hemispheric Differences in the Recognition of Environmental Sounds

ABSTRACT— Recent work has found support for two dissociable and parallel neural subsystems underlying object and shape recognition in the visual domain: an abstract-category subsystem that operates more effectively in the left cerebral hemisphere than in the right, and a specific-exemplar subsystem that operates more effectively in the right hemisphere than in the left. Evidence of this asymmetry has been observed for linguistic stimuli (words, pseudoword forms) and nonlinguistic stimuli (objects). In the auditory domain, we previously found hemispheric asymmetries in priming effects using linguistic stimuli (spoken words). In the present study, we conducted four long-term repetition-priming experiments to investigate whether such hemispheric asymmetries would be observed for nonlinguistic auditory stimuli (environmental sounds) as well. The results support the dissociable-subsystems theory. Specificity effects were obtained when sounds were presented to the left ear (right hemisphere), but not when sounds were presented to the right ear (left hemisphere). Theoretical implications are discussed.     

The perception of stress and lateralization of prosody

Three separate dichotic listening tasks were run to determine ear superiority for stress identification. When subjects were asked to identify stress placement in real word minimal stress pairs (hótdog vs. hot dóg), they demonstrated a right ear superiority. When these tokens were filtered so that phonetic and semantic information was eliminated and only the stress pattern remained, a different group of subjects showed a left ear advantage. Finally, with nonsense word counterparts to word stress pairs (bótgog vs. bot góg) preserving phonetic information but lacking semantic content, no ear asymmetry was found. These results suggest that as the linguistic significance of the stimuli is reduced, thereby lessening the linguistic function of stress, there is a less dominant involvement of the left hemisphere in stress processing. Results are discussed in relation to a theory of a functional integration of prosodic and segmental speech components that is paralleled by a working partnership of left and right hemisphere.     

Phonological processing of words in right- and left-handers

It is commonly accepted that phonology is the exclusive domain of the left hemisphere. However, this pattern of lateralization, which posits a right visual field advantage, has been questioned by several studies. In fact, certain factors such as characteristics of the stimuli and subjects' handedness can modulate the right visual field advantage. Thus, the goal of this study was to compare the hemispheric dynamics of right-handers and left-handers during a divided visual field presentation of words that varied in terms of their phonological transparency. For non-transparent words, the left hemisphere seems more competent in both handedness groups. With regard to transparent words, the right hemisphere of both groups also appears competent. Surprisingly, left-handers achieved optimal processing with a functionally isolated left hemisphere, whereas right-handers needed the participation of both hemispheres. The pattern of performance cannot be fully explained by either the callosal or the direct access model.     

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

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

Ear asymmetries on a selective attentional task

To ascertain whether there are ear-hemisphere asymmetries of selective attention, signal stimuli (tonal sequences) were presented monaurally with and without complex maskers (music and speech). The right ear-left hemisphere was more disrupted by language maskers; the left ear-right hemisphere was more disrupted by music maskers. These results suggest that there are hemispheric asymmetries of selective attention and that the ear hemisphere that usually processes a class of stimuli has greater difficulty filtering out those stimuli than does the nonspecialized hemisphere.     

Cross-modal matching and association under sodium amytal

Anatomical and theoretical considerations, as well as experimental findings, have yielded conflicting points of view regarding the abilities of the right hemisphere in man to accomplish cross-modal transfer of information. Auditory—visual cross-modal matching (CMM) and association (CMA) abilities of the left and right hemisphere (LH and RH) were tested, using the Wada intracarotid sodium amytal technique. It was found that the RH performed slightly better on these tasks than the LH. These findings contrast with results of other techniques which indicate that the RH cannot perform CMM and CMA.     

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.     

Hemispheric asymmetries for gap detection depend on noise type

Studies of temporal processing asymmetries in the auditory modality have not produced consistent results. Some investigators have found a left hemisphere advantage, but others have failed to replicate this result. The present experiment investigated the possibility that differing properties of the noise employed between these experiments could be responsible for the conflicting results. Short bursts (300 ms) of white or brown noise were delivered monaurally to 42 participants. Half of the stimuli contained 3-5 ms gaps of silence. A right ear (left hemisphere) advantage in accuracy was observed in the white noise condition, but there was no such difference in the brown noise condition. This result demonstrates that the different acoustic properties of the stimuli between experiments can account for some of the discrepancies in their findings, and supports the position that the left hemisphere is superior at processing rapid temporal changes.     

The effects of simulated stuttering and prolonged speech on the neural activation patterns of stuttering and nonstuttering adults

Functional magnetic resonance imaging was used to investigate the neural correlates of passive listening, habitual speech and two modified speech patterns (simulated stuttering and prolonged speech) in stuttering and nonstuttering adults. Within-group comparisons revealed increased right hemisphere biased activation of speech-related regions during the simulated stuttered and prolonged speech tasks, relative to the habitual speech task, in the stuttering group. No significant activation differences were observed within the nonstuttering participants during these speech conditions. Between-group comparisons revealed less left superior temporal gyrus activation in stutterers during habitual speech and increased right inferior frontal gyrus activation during simulated stuttering relative to nonstutterers. Stutterers were also found to have increased activation in the left middle and superior temporal gyri and right insula, primary motor cortex and supplementary motor cortex during the passive listening condition relative to nonstutterers. The results provide further evidence for the presence of functional deficiencies underlying auditory processing, motor planning and execution in people who stutter, with these differences being affected by speech manner.     

Lateralization of auditory rhythm length in temporal lobe lesions

In the visual modality, short rhythmic stimuli have been proven to be better processed (sequentially) by the left hemisphere, while longer rhythms appear to be better (holistically) processed by the right hemisphere. This study was set up to see if the same holds in the auditory modality. The rhythm task as originally designed by Seashore was computerized and is part of the Fepsy Neuropsychological battery. This task was performed by 85 patients with intractable temporal lobe epilepsy (left TLE = 32; right TLE = 53) enrolled in the Dutch Collaborative Epilepsy Surgery Program. They performed the task before and 6 months after surgery. The task consists of 30 pairs of rhythmic patterns in 3 series of 10 items. The series contains patterns of 5, 6, or 7 notes. The purpose is to indicate whether the two patterns are the same or different. Reaction times are also measured. If the hypothesis is true, the short-item sequence will be better processed by patients with right temporal lobe epilepsy (nonimpaired left temporal lobe), the longer sequence will be better processed by the left temporal epilepsy group (nonimpaired right temporal lobe). No overall laterality effect on rhythm perception could be found and no difference was found between both test moments. IQ did not correlate with rhythm performance. However, there was an interaction effect of laterality and rhythm length on performance and reaction time. This effect can be explained by the increase after the operation of the score of the left focus group and a decrease in the right focus group on the longer rhythms. This effect was somewhat less strong in the reaction times: a clear tendency for faster reaction times after surgery in the left and longer reaction times in the right focus group. The effect could not be explained for by the difference in extent of resection in either temporal lobe. This study showed that memory for and discrimination of auditory rhythm is dependent on which hemisphere is used in processing. The effect could be demonstrated for the right hemisphere, which uses a holistic processing of stimuli, which outperforms the left in rhythms consisting of a long sequence. In left temporal resections an improvement occurs on the longer rhythms and in right temporal resections the performance on the longest rhythms decreases.     

Pure word deafness and the bilateral processing of the speech code

The analysis of pure word deafness (PWD) suggests that speech perception, construed as the integration of acoustic information to yield representations that enter into the linguistic computational system, (i) is separable in a modular sense from other aspects of auditory cognition and (ii) is mediated by the posterior superior temporal cortex in both hemispheres. PWD data are consistent with neuropsychological and neuroimaging evidence in a manner that suggests that the speech code is analyzed bilaterally. The typical lateralization associated with language processing is a property of the computational system that acts beyond the analysis of the input signal. The hypothesis of the bilateral mediation of the speech code does not imply that both sides execute the same computation. It is proposed that the speech signal is asymmetrically analyzed in the time domain, with left‐hemisphere mechanisms preferentially extracting information over shorter (25–50 ms) temporal integration windows and right mechanisms over longer (150–250 ms) windows.     

Colateralization of Broca's area and the visual word form area in left-handers: fMRI evidence

Language production has been found to be lateralized in the left hemisphere (LH) for 95% of right-handed people and about 75% of left-handers. The prevalence of atypical right hemispheric (RH) or bilateral lateralization for reading and colateralization of production with word reading laterality has never been tested in a large sample. In this study, we scanned 57 left-handers who had previously been identified as being clearly left (N = 30), bilateral (N = 7) or clearly right (N = 20) dominant for speech on the basis of fMRI activity in the inferior frontal gyrus (pars opercularis/pars triangularis) during a silent word generation task. They were asked to perform a lexical decision task, in which words were contrasted against checkerboards, to test the lateralization of reading in the ventral occipitotemporal region. Lateralization indices for both tasks correlated significantly (r = 0.59). The majority of subjects showed most activity during lexical decision in the hemisphere that was identified as their word production dominant hemisphere. However, more than half of the sample (N = 31) had bilateral activity for the lexical decision task without a clear dominant role for either the LH or RH, and three showed a crossed frontotemporal lateralization pattern. These findings have consequences for neurobiological models relating phonological and orthographic processes, and for lateralization measurements for clinical purposes.