Left Hemisphere Specialization For Language Research Articles

Saccade accuracy as an indicator of the competition between functional asymmetries in vision.

Hemispheric specialization refers to the fact that cerebral hemispheres are not equivalent and that cognitive processes are lateralized in the brain. Although the potential links between handedness and the left hemisphere specialization for language have been widely studied, little attention has been paid to other motor preferences, such as eye dominance, that also are lateralized in the brain. For example, saccadic accuracy is higher in the hemifield contralateral to the dominant eye compared to the ipsilateral hemifield. Saccade accuracy is, however, also known to be sensitive to other functional asymmetries, such as the lateralization of visuo-spatial attention in the right hemisphere of the brain. Using a global effect paradigm in three different saccade latency ranges, we here propose to use saccade accuracy as an indicator of visual functional asymmetries. We show that for the shortest latencies, saccade accuracy is higher in the left than in the right visual hemifield, which could be due to the lateralization of visuo-spatial attention in the right hemisphere. For the longest latencies, however, saccade accuracy is higher toward the right than the left hemifield, probably due to the lateralization of local and global processing in the left and right hemispheres, respectively. These results could have a major impact on studies designed to measure the degree of lateralization of individuals. We here discuss both the theoretical and clinical contributions of these results.

Early asymmetric inter-hemispheric transfer in the auditory network: insights from infants with corpus callosum agenesis.

The left hemisphere specialization for language is a well-established asymmetry in the human brain. Structural and functional asymmetries are observed as early as the prenatal period suggesting genetically determined differences between both hemispheres. The corpus callosum is a large tract connecting mostly homologous areas; some have proposed that it might participate in an enhancement of the left-hemispheric advantage to process speech. To investigate its role in early development, we compared 13 3-4-month-old infants with an agenesis of the corpus callosum ("AgCC") with 18 typical infants using high-density electroencephalography in an auditory task. We recorded event-related potentials for speech stimuli (syllables and babbling noise), presented binaurally (same syllable in both ears), monaurally (babbling noise in one ear) and dichotically (syllable in one ear and babbling noise in the other ear). In response to these stimuli, both groups developed an anterior positivity synchronous with a posterior negativity, yet the topography significantly differed between groups likely due to the atypical gyration of the medial surface in AgCC. In particular, the anterior positivity was lateral in AgCC infants while it covered the midline in typical infants. We then measured the latencies of the main auditory response (P2 at this age) for the different conditions on the symmetrical left and right clusters. The main difference between groups was a ~ 60ms delay in typical infants relative to AgCC, for the ipsilateral response (i.e. left hemisphere) to babbling noise presented in the left ear, whereas no difference was observed in the case of right-ear stimulation. We suggest that our results highlight an asymmetrical callosal connectivity favoring the right-to-left hemisphere direction in typical infants. This asymmetry, similar to recent descriptions in adults, might contribute to an enhancement of left lateralization for language processing beyond the initial cortical left-hemisphere advantage.

A multifactorial investigation of captive gorillas’ intraspecific gestural laterality

ABSTRACTMultifactorial investigations of intraspecific laterality of primates’ gestural communication aim to shed light on factors that underlie the evolutionary origins of human handedness and language. This study assesses gorillas’ intraspecific gestural laterality considering the effect of various factors related to gestural characteristics, interactional context and sociodemographic characteristics of signaller and recipient. Our question was: which factors influence gorillas’ gestural laterality? We studied laterality in three captive groups of gorillas (N = 35) focusing on their most frequent gesture types (N = 16). We show that signallers used predominantly their hand ipsilateral to the recipient for tactile and visual gestures, whatever the emotional context, gesture duration, recipient’s sex or the kin relationship between both interactants, and whether or not a communication tool was used. Signallers’ contralateral hand was not preferentially used in any situation. Signallers’ right-hand use was more pronounced in negative contexts, in short gestures, when signallers were females and its use increased with age. Our findings showed that gorillas’ gestural laterality could be influenced by different types of social pressures thus supporting the theory of the evolution of laterality at the population level. Our study also evidenced that some particular gesture categories are better markers than others of the left-hemisphere language specialization.

Semantics is crucial for the right-hemisphere involvement in metaphor processing: Evidence from mouth asymmetry during speaking

Research on the neural basis of metaphor provides contradicting evidence about the role of right and left hemispheres. We used the mouth-opening asymmetry technique to investigate the relative involvement of the two hemispheres whilst right-handed healthy male participants explained the meaning of English phrases. This technique is based on the contralateral cortical control of the facial musculature and reflects the relative hemispheric involvement during different cognitive tasks. In particular, right-handers show a right-sided mouth asymmetry (right side of the mouth opens wider than the left) during linguistic tasks, thus reflecting the left-hemisphere specialization for language. In the current study, we compared the right-sided mouth asymmetry during metaphor explanation (e.g., explain the meaning of the phrase “to spin a yarn”) and concrete explanation (e.g., explain the meaning of the phrase “to spin a golf ball”) and during the production of content and function words. The expected right-sided mouth asymmetry reduced during metaphorical compared to concrete explanations suggesting the relative right-hemispheric involvement for metaphor processing. Crucially, this right-sided mouth asymmetry reduction was particularly pronounced for the production of content words. Thus, we concluded that semantics is crucial to the right-hemispheric involvement for metaphorical speech production.

Handedness for grasping objects and declarative pointing: A longitudinal study

It is still unclear whether infants become right-handed because of their left-hemisphere specialization for language (through gestural communication for instance), whether they speak predominantly with their left hemisphere because of this hemisphere's superiority in controlling sequential actions which first results in right-handedness, or whether the two lateralization processes develop independently. To tackle this question, we followed 26 human infants from 8 to 20 months to evaluate the temporal relationship between the emergence of hand preference for grasping objects and for declarative pointing (communicative gesture). Our results show that when grasping and pointing are compared in similar conditions, with objects presented in several spatial positions, the tendency to use the right hand is significantly larger for pointing than for grasping, and both hand preferences are loosely correlated. This suggests that, at least at the age studied here, hand preferences for grasping and for declarative pointing develop relatively independently.

Effect of Familial Sinistrality on Planum Temporale Surface and Brain Tissue Asymmetries

The impact of having left-handers (LHs) among one's close relatives, called familial sinistrality (FS), on neuroanatomical markers of left-hemisphere language specialization was studied in 274 normal adults, including 199 men and 75 women, among whom 77 men and 27 women were positive for FS. Measurements of the surface of a phonological cortical area, the "planum temporale" (PT), and gray and white matter hemispheric volumes and asymmetries were made using brain magnetic resonance images. The size of the left PT of subjects with left-handed close relatives (FS+) was reduced by 10%, decreasing with the number of left-handed relatives, and lowest when the subject's mother was left-handed. Such findings had no counterparts in the right hemisphere, and the subject's handedness and sex were found to have no significant effect or interaction with FS on the left PT size. The FS+ subjects also exhibited increased gray matter volume, reduced hemispheric gray matter leftward asymmetry, and, in LHs, reduced strength of hand preference. These results add to the increasing body of evidence suggesting multiple and somewhat independent mechanisms for the inheritance of hand and language lateralization.

Brain, language, and handedness: a family affair

AbstractThe left planum temporale is a marker of left hemisphere language specialization. We investigated the effect of individual handedness and familial sinistrality on left planum temporale surface area and found the size is reduced in proportion with the number of left-handed immediate family members and is lowest when one's mother is left-handed. This reduction is independent of an individual's handedness or sex and has no counterpart in the right hemisphere.

Neuroanatomical Correlates of Handedness for Tool Use in Chimpanzees (Pan Troglodytes)

It has been hypothesized that cognitive mechanisms underlying lateralized complex motor actions associated with tool use in chimpanzees may have set the stage for the evolution of left-hemisphere specialization for language and speech in humans. Here we report evidence that asymmetries in the homologues to Broca's and Wernicke's areas are associated with handedness for tool use in chimpanzees. These results suggest that the neural substrates of tool use may have served as a preadaptation for the evolution of language and speech in modern humans.

Hemispheric asymmetries in language-related pathways: A combined functional MRI and tractography study

Functional lateralization is a feature of human brain function, most apparent in the typical left-hemisphere specialization for language. A number of anatomical and imaging studies have examined whether structural asymmetries underlie this functional lateralization. We combined functional MRI (fMRI) and diffusion-weighted imaging (DWI) with tractography to study 10 healthy right-handed subjects. Three language fMRI paradigms were used to define language-related regions in inferior frontal and superior temporal regions. A probabilistic tractography technique was then employed to delineate the connections of these functionally defined regions. We demonstrated consistent connections between Broca's and Wernicke's areas along the superior longitudinal fasciculus bilaterally but more extensive fronto-temporal connectivity on the left than the right. Both tract volumes and mean fractional anisotropy (FA) were significantly greater on the left than the right. We also demonstrated a correlation between measures of structure and function, with subjects with more lateralized fMRI activation having a more highly lateralized mean FA of their connections. These structural asymmetries are in keeping with the lateralization of language function and indicate the major structural connections underlying this function.

Asymmetries for the visual expression and perception of speech

This study explored asymmetries for movement, expression and perception of visual speech. Sixteen dextral models were videoed as they articulated: ‘bat,’ ‘cat,’ ‘fat,’ and ‘sat.’ Measurements revealed that the right side of the mouth was opened wider and for a longer period than the left. The asymmetry was accentuated at the beginning and ends of the vocalization and was attenuated for words where the lips did not articulate the first consonant. To measure asymmetries in expressivity, 20 dextral observers watched silent videos and reported what was said. The model’s mouth was covered so that the left, right or both sides were visible. Fewer errors were made when the right mouth was visible compared to the left—suggesting that the right side is more visually expressive of speech. Investigation of asymmetries in perception using mirror-reversed clips revealed that participants did not preferentially attend to one side of the speaker’s face. A correlational analysis revealed an association between movement and expressivity whereby a more motile right mouth led to stronger visual expressivity of the right mouth. The asymmetries are most likely driven by left hemisphere specialization for language, which causes a rightward motoric bias.

The Distribution and Development of Handedness for Manual Gestures in Captive Chimpanzees (Pan troglodytes)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, non-communicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.

Atypical hemispheric specialization for language in right-handed schizophrenia patients

The literature suggests that schizophrenia could be related to a failure in the setting up of left hemisphere dominance for language. We sought to determine hemispheric specialization for language in schizophrenic patients, using functional magnetic resonance imaging. Twenty-one right-handed patients with DSM-IV schizophrenia and 21 right-handed control subjects matched by age, gender, and level of education were recruited. Fractional blood oxygen level dependent (BOLD) signal variations in anatomic regions of interest were compared between groups. Functional asymmetry indices (FAIs) were calculated in a region (LANG) resulting from the merging of activated regions showing a Group x Hemisphere interaction. The FAI difference between each patient and their matched control subject was computed. We found lower BOLD signal changes in patients as compared with their control subjects in a network comprising areas of the left middle temporal gyrus, the left angular gyrus, and the pars triangularis of the left inferior frontal gyrus, merged to constitute LANG. The intra-pair differences of FAIs in this area showed that 76% of the patients exhibited less leftward functional asymmetry than their matched control subjects, including six patients with a rightward asymmetry. These results demonstrated the existence of an anomaly in left hemisphere specialization for language in schizophrenic subjects.

Hemispheric Specialization for Language in Children with Different Types of Specific Language Impairment

The aim of the study was to investigate whether children with Specific Language Impairment (SLI) show reduced left hemisphere specialization for language and, if so, whether it is associated with a deficit in phonological encoding and a specific type of SLI (Mixed Receptive-Expressive, Expressive, Phonological). We adopted two dichotic listening paradigms, which differed in the phonological similarity of the presented words (Fused and Non-Fused dichotic words tests), as well as a phonological working memory test. Participants included 34 pre-school and school age children affected by SLI. On the dichotic tests, as a group the children with SLI showed a reduced pattern of left hemisphere specialization for language compared to age-matched normal children, with significant differences only in the Fused condition. However, the pattern of hemispheric specialization varied depending on the type of SLI, with reduced left hemisphere specialization in the Expressive type and, to a lesser extent, in the Phonological type of SLI, but not in the Receptive-Expressive type. The three subgroups also differed in phonological processing abilities and the incidence of a positive family history for language disorders: the Receptive-Expressive group performed worse on the working memory and dichotic tests and the Expressive and Phonological groups presented high frequency for familial language disorder. These results suggest that different subtypes of SLI are not different manifestations of the same underlying disorder, but represent pathological conditions that have distinct markers both at the behavioral and neurofunctional level.

Planum temporale asymmetry and models of dominance for language: a reappraisal.

A classical developmental model for the left hemisphere specialization for language has been proposed based on the observation of a positive correlation coefficient between the value of an index of asymmetry between the left and right planum temporale surface areas and the size of the right planum temporale. Here, we demonstrate that such correlation is a mathematical artefact and thus should not serve as a basis for establishing models of language hemispheric dominance.

Handedness in Chimpanzees (Pan troglodytes) Is Associated With Asymmetries of the Primary Motor Cortex but Not With Homologous Language Areas.

The neurobiology of hand preferences in nonhuman primates is poorly understood. In this study, the authors report the first evidence of an association between hand preference and precentral gyrus-morphology in chimpanzees (Pan troglodytes). Hand preferences did not significantly correlate with other asymmetric brain regions associated with language functions in humans including the planum temporale and frontal operculum. The overall results suggest that homologous regions of the motor cortex control hand preferences in humans and apes and that these functions evolved independently of left-hemisphere specialization for language and speech.

The Lateral Asymmetry of the Human Brain Studied by Volumetric Magnetic Resonance Imaging

Improvements in in vivo imaging methods have boosted research on brain asymmetry aimed at further establishing putative anatomical substrates for brain functional lateralization and particularly to explain left-hemisphere specialization for language. We analyzed volume asymmetries for major anatomical divisions of the lateral (perisylvian) brain region and their relative white matter content. A total of 100 healthy right-handed subjects were examined with 3D magnetic resonance imaging (MRI). The insular plane was used to limit the lateral brain, and the sylvian fissure and central sulcus to define frontal, parietal, temporal, and temporo-parieto-occipital regions. Results revealed a frontal region showing similar volumes in both hemispheres, a parietal region and a temporal region both larger in the left hemisphere, and a temporo-parieto-occipital region with predominantly right-sided asymmetry. Volume measurements of the parietal, temporal, and temporo-parieto-occipital regions complemented each other and accounted for 58% of planum temporale area variations. All study regions showed significant asymmetry for relative white matter content (percentage of white matter relative to region volume). White matter asymmetry, however, was particularly relevant for the frontal and temporal regions showing a highly frequent left-sided pattern (frontal region, 90%; temporal region, 91% of subjects). Leftward asymmetry in these two regions occurred in both genders, although hemisphere differences were significantly larger in men. Results from this MRI volume analysis of structural asymmetries in the lateral brain region complement data obtained by other methods and suggest a high occurrence of leftward asymmetry for relative white matter content in language-related regions.

The Lateral Asymmetry of the Human Brain Studied by Volumetric Magnetic Resonance Imaging

Improvements in in vivo imaging methods have boosted research on brain asymmetry aimed at further establishing putative anatomical substrates for brain functional lateralization and particularly to explain left-hemisphere specialization for language. We analyzed volume asymmetries for major anatomical divisions of the lateral (perisylvian) brain region and their relative white matter content. A total of 100 healthy right-handed subjects were examined with 3D magnetic resonance imaging (MRI). The insular plane was used to limit the lateral brain, and the sylvian fissure and central sulcus to define frontal, parietal, temporal, and temporo-parieto-occipital regions. Results revealed a frontal region showing similar volumes in both hemispheres, a parietal region and a temporal region both larger in the left hemisphere, and a temporo-parieto-occipital region with predominantly right-sided asymmetry. Volume measurements of the parietal, temporal, and temporo-parieto-occipital regions complemented each other and accounted for 58% of planum temporale area variations. All study regions showed significant asymmetry for relative white matter content (percentage of white matter relative to region volume). White matter asymmetry, however, was particularly relevant for the frontal and temporal regions showing a highly frequent left-sided pattern (frontal region, 90%; temporal region, 91% of subjects). Leftward asymmetry in these two regions occurred in both genders, although hemisphere differences were significantly larger in men. Results from this MRI volume analysis of structural asymmetries in the lateral brain region complement data obtained by other methods and suggest a high occurrence of leftward asymmetry for relative white matter content in language-related regions.

Asymmetries in preparation for action

The origins and nature of hemispheric specialization of action control are unclear. A review of some recent evidence suggests that the right hemisphere interprets spatial relationships whereas the left deals with temporal control of movement. Contrary to the popular view, specialization of the right hemisphere for spatial representations might have preceded left hemisphere specialization for language and movement.

Functional lateralization for auditory temporal processing in male and female rats.

In 3 studies adult male rats showed significantly better discrimination of tone sequences with the right ear than with the left ear. This result parallels similar findings of left-hemisphere specialization for auditory temporal processing in humans and nonhuman primates. Furthermore, because clinical evidence supports a link between specialization of the left hemisphere for auditory temporal processing and for speech processing, these results may reflect evolutionary precursors to left-hemisphere language specialization. Because male rats showed a stronger ear advantage than female rats, the findings may relate to evidence of a stronger right ear advantage in men than in women. Finally, results suggest that neonatal handling enhances lateralization for auditory temporal processing in both sexes. Combined results implicate neuroendocrine mechanisms as important factors in the development of lateralization for auditory temporal processing.

Left-Hemisphere Language Specialization: Consistency in Hand Preference and Sex Differences

KEE, DANIEL W.; GOTTFRIED, ALLEN W.; BATHURST, KAY; and BROWN, KATHLEEN. Left-Hemisphere Language Specialization: Consistency in Hand Preference and Sex Differences. CHILD DEVELOPMENT, 1987, 58, 718-724. A dual task/time-sharing procedure was used to evaluate left-hemisphere language specialization in right-handed children tested at ages 5 and 6. These children from a longitudinal study had been previously classified as consistent versus nonconsistent in their hand preference measured every 6 months between 18 and 42 months of age. Results showed that both male hand-preference groups demonstrated asymmetric interference in dual task tapping performance indicative of left-hemisphere language specialization. In contrast, only females associated with consistency in hand preference across time showed this asymmetry.