Positional and surface area asymmetry of the human cerebral cortex

Previous studies of cortical asymmetry have relied mainly on voxel-based morphometry (VBM), or manual segmentation of regions of interest. This study uses fully automated, surface-based techniques to analyse position and surface area asymmetry for the mid-surfaces of 112 right-handed subjects' cortical hemispheres from a cohort of young adults. Native space measurements of local surface area asymmetry and vertex position asymmetry were calculated from surfaces registered to a previously validated hemisphere-unbiased surface-based template. Our analysis confirms previously identified hemispheric asymmetries (Yakovlevian torque, frontal and occipital petalia) in enhanced detail. It does not support previous findings of gender/asymmetry interactions or rightward planum parietale areal increase. It reveals several new findings, including a striking leftward increase in surface area of the supramarginal gyrus (peak effect 18%), compared with a smaller areal increase in the left Heschl's gyrus and planum temporale region (peak effect 8%). A second finding was rightward increase in surface area (peak effect 10%) in a band around the medial junction between the occipital lobe, and parietal and temporal lobes. By clearly separating out the effects of structural translocation and surface area change from those of thickness and curvature, this study resolves the confound of these variables inherent in VBM studies.     

Cerebral asymmetry and the effects of sex and handedness on brain structure: a voxel-based morphometric analysis of 465 normal adult human brains

We used voxel-based morphometry (VBM) to examine human brain asymmetry and the effects of sex and handedness on brain structure in 465 normal adults. We observed significant asymmetry of cerebral grey and white matter in the occipital, frontal, and temporal lobes (petalia), including Heschl's gyrus, planum temporale (PT) and the hippocampal formation. Males demonstrated increased leftward asymmetry within Heschl's gyrus and PT compared to females. There was no significant interaction between asymmetry and handedness and no main effect of handedness. There was a significant main effect of sex on brain morphology, even after accounting for the larger global volumes of grey and white matter in males. Females had increased grey matter volume adjacent to the depths of both central sulci and the left superior temporal sulcus, in right Heschl's gyrus and PT, in right inferior frontal and frontomarginal gyri and in the cingulate gyrus. Females had significantly increased grey matter concentration extensively and relatively symmetrically in the cortical mantle, parahippocampal gyri, and in the banks of the cingulate and calcarine sulci. Males had increased grey matter volume bilaterally in the mesial temporal lobes, entorhinal and perirhinal cortex, and in the anterior lobes of the cerebellum, but no regions of increased grey matter concentration.     

Mapping an intrinsic MR property of gray matter in auditory cortex of living humans: a possible marker for primary cortex and hemispheric differences

Recently, magnetic resonance properties of cerebral gray matter have been spatially mapped--in vivo--over the cortical surface. In one of the first neuroscientific applications of this approach, this study explores what can be learned about auditory cortex in living humans by mapping longitudinal relaxation rate (R1), a property related to myelin content. Gray matter R1 (and thickness) showed repeatable trends, including the following: (1) Regions of high R1 were always found overlapping posteromedial Heschl's gyrus. They also sometimes occurred in planum temporale and never in other parts of the superior temporal lobe. We hypothesize that the high R1 overlapping Heschl's gyrus (which likely indicates dense gray matter myelination) reflects auditory koniocortex (i.e., primary cortex), a heavily myelinated area that shows comparable overlap with the gyrus. High R1 overlapping Heschl's gyrus was identified in every instance suggesting that R1 may ultimately provide a marker for koniocortex in individuals. Such a marker would be significant for auditory neuroimaging, which has no standard means (anatomic or physiologic) for localizing cortical areas in individual subjects. (2) Inter-hemispheric comparisons revealed greater R1 on the left on Heschl's gyrus, planum temporale, superior temporal gyrus and superior temporal sulcus. This asymmetry suggests greater gray matter myelination in left auditory cortex, which may be a substrate for the left hemisphere's specialized processing of speech, language, and rapid acoustic changes. These results indicate that in vivo R1 mapping can provide new insights into the structure of human cortical gray matter and its relation to function.     

Handedness and cerebral anatomical asymmetries in young adult males

Using voxel-based morphometry, we measured the cerebral anatomical asymmetries in a sample of 56 young right-handed males and then compared voxelwise asymmetry indices of these subjects to those of 56 young left-handed males. In the right-handed, the clusters of grey matter asymmetry corresponding to the leftward occipital petalia and planum temporale asymmetries were retrieved. Strong rightward temporo-parietal asymmetries were also observed, but the rightward grey matter asymmetry in the frontal lobe was less massive than previously described. Group comparisons of left- and right-handed subjects' asymmetry maps, performed at a statistical threshold not corrected for multiple comparisons, revealed significant effects of handedness on this pattern of anatomical asymmetry in frontal regions, notably in the lower central and precentral sulci, and also in the planum temporale, with right-handed subjects being more leftward asymmetric. Concerning white matter, although almost no focal differences between left- and right-handed subjects were detected, volumetric analyses at the hemispheric level revealed a leftward asymmetry, which happened to be significantly less marked in the left-handed. This latter result, together with the pattern of leftward white matter asymmetries, suggested that anatomical correlates of the left hemispheric specialization for language would exist in white matter. In the population we studied, differences in anatomical asymmetry between left- and right-handed subjects provided structural arguments for a greater functional ambilaterality in left-handed subjects.     

Neural circuitry underlying sentence-level linguistic prosody

This study investigates the neural substrates underlying the perception of two sentence-level prosodic phenomena in Mandarin Chinese: contrastive stress (initial vs. final emphasis position) and intonation (declarative vs. interrogative modality). In an fMRI experiment, Chinese and English listeners were asked to selectively attend to either stress or intonation in paired 3-word sentences, and make speeded-response discrimination judgments. Between-group comparisons revealed that the Chinese group exhibited significantly greater activity in the left supramarginal gyrus and posterior middle temporal gyrus relative to the English group for both tasks. These same two regions showed a leftward asymmetry in the stress task for the Chinese group only. For both language groups, rightward asymmetries were observed in the middle portion of the middle frontal gyrus across tasks. All task effects involved greater activity for the stress task as compared to intonation. A left-sided task effect was observed in the posterior middle temporal gyrus for the Chinese group only. Both language groups exhibited a task effect bilaterally in the intraparietal sulcus. These findings support the emerging view that speech prosody perception involves a dynamic interplay among widely distributed regions not only within a single hemisphere but also between the two hemispheres. This model of speech prosody processing emphasizes the role of right hemisphere regions for complex-sound analysis, whereas task-dependent regions in the left hemisphere predominate when language processing is required.     

Representation of lateralization and tonotopy in primary versus secondary human auditory cortex

Functional MRI was performed to investigate differences in the basic functional organization of the primary and secondary auditory cortex regarding preferred stimulus lateralization and frequency. A modified sparse acquisition scheme was used to spatially map the characteristics of the auditory cortex at the level of individual voxels. In the regions of Heschl's gyrus and sulcus that correspond with the primary auditory cortex, activation was systematically strongest in response to contralateral stimulation. Contrarily, in the surrounding secondary active regions including the planum polare and the planum temporale, large-scale preferences with respect to stimulus lateralization were absent. Regarding optimal stimulus frequency, low- to high-frequency spatial gradients were discernable along the Heschl's gyrus and sulcus in anterolateral to posteromedial direction, especially in the right hemisphere, consistent with the presence of a tonotopic organization in these primary areas. However, in the surrounding activated secondary areas frequency preferences were erratic. Lateralization preferences did not depend on stimulus frequency, and frequency preferences did not depend on stimulus lateralization. While the primary auditory cortex is topographically organized with respect to physical stimulus properties (i.e., lateralization and frequency), such organizational principles are no longer obvious in secondary and higher areas. This suggests a neural re-encoding of sound signals in the transition from primary to secondary areas, possibly in relation to auditory scene analysis and the processing of auditory objects.     

Effective connectivity analysis demonstrates involvement of premotor cortex during speech perception

Several reports of premotor cortex involvement in speech perception have been put forward. Still, the functional role of premotor cortex is under debate. In order to investigate the functional role of premotor cortex, we presented parametrically varied speech stimuli in both a behavioral and functional magnetic resonance imaging (fMRI) study. White noise was transformed over seven distinct steps into a speech sound and presented to the participants in a randomized order. As control condition served the same transformation from white noise into a music instrument sound. The fMRI data were modelled with Dynamic Causal Modeling (DCM) where the effective connectivity between Heschl's gyrus, planum temporale, superior temporal sulcus and premotor cortex were tested. The fMRI results revealed a graded increase in activation in the left superior temporal sulcus. Premotor cortex activity was only present at an intermediate step when the speech sounds became identifiable but were still distorted but was not present when the speech sounds were clearly perceivable. A Bayesian model selection procedure favored a model that contained significant interconnections between Heschl's gyrus, planum temporal, and superior temporal sulcus when processing speech sounds. In addition, bidirectional connections between premotor cortex and superior temporal sulcus and from planum temporale to premotor cortex were significant. Processing non-speech sounds initiated no significant connections to premotor cortex. Since the highest level of motor activity was observed only when processing identifiable sounds with incomplete phonological information, it is concluded that premotor cortex is not generally necessary for speech perception but may facilitate interpreting a sound as speech when the acoustic input is sparse.     

Asymmetry in the Human Motor Cortex and Handedness

Handedness is one of the most obvious functional asymmetries, but its relation to an anatomical asymmetry of the hand representation area in the motor cortex has not been demonstrated. This would be a crucial test for the hypothesis of structure–function correlation in cortical motor organization. Using magnetic resonance morphometry, we show for the first time that the depth of the central sulcus is related to handedness. In right-handers, the left central sulcus is deeper than the right, and vice versa in left-handers. Macrostructural asymmetry is complemented by a microstructural left-larger-than-right asymmetry in neuropil volume (i.e., tissue compartment containing dendrites, axons, and synapses) in Brodmann's area 4. These asymmetries suggest that hand preference is associated with increased connectivity (demonstrated by an increased neuropil compartment in left area 4) and an increased intrasulcal surface of the precentral gyrus in the dominant hemisphere.     

Automatic analysis of cerebral asymmetry: an exploratory study of the relationship between brain torque and planum temporale asymmetry

Leftward occipital and rightward frontal lobe asymmetry (brain torque) and leftward planum temporale asymmetry have been consistently reported in postmortem and in vivo neuroimaging studies of the human brain. Here automatic image analysis techniques are applied to quantify global and local asymmetries, and investigate the relationship between brain torque and planum temporale asymmetries on T1-weighted magnetic resonance (MR) images of 30 right-handed young healthy subjects (15 male, 15 female). Previously described automatic cerebral hemisphere extraction and 3D interhemispheric reflection-based methods for studying brain asymmetry are applied with a new technique, LowD (Low Dimension), which enables automatic quantification of brain torque. LowD integrates extracted left and right cerebral hemispheres in columns orthogonal to the midsagittal plane (2D column maps), and subsequently integrates slices along the brain's anterior-posterior axis (1D slice profiles). A torque index defined as the magnitude of occipital and frontal lobe asymmetry is computed allowing exploratory investigation of relationships between this global asymmetry and local asymmetries found in the planum temporale. LowD detected significant torque in the 30 subjects with occipital and frontal components found to be highly correlated (P<0.02). Significant leftward planum temporale asymmetry was detected (P<0.05), and the torque index correlated with planum temporale asymmetry (P<0.001). However, torque and total brain volume were not correlated. Therefore, although components of cerebral asymmetry may be related, their magnitude is not influenced by total hemisphere volume. LowD provides increased sensitivity for detection and quantification of brain torque on an individual subject basis, and future studies will apply these techniques to investigate the relationship between cerebral asymmetry and functional laterality.     

The morphometry of auditory cortex in the congenitally deaf measured using MRI

The study of congenitally deaf individuals provides a unique opportunity to understand the organization and potential for reorganization of human auditory cortex. We used magnetic resonance imaging (MRI) to examine the structural organization of two auditory cortical regions, Heschl's gyrus (HG) and the planum temporale (PT), in deaf and hearing subjects. The results show preservation of cortical volume in HG and PT of deaf subjects deprived of auditory input since birth. Measurements of grey and white matter, as well as the location and extent of these regions in the deaf showed complete overlap both with matched controls and with previous samples of hearing subjects. The results of the manual volume measures were supported by findings from voxel-based morphometry analyses that showed increased grey-matter density in the left motor hand area of the deaf, but no differences between the groups in any auditory cortical region. This increased cortical density in motor cortex may be related to more active use of the dominant hand in signed languages. Most importantly, expected interhemispheric asymmetries in HG and PT thought to be related to auditory language processing were preserved in these deaf subjects. These findings suggest a strong genetic component in the development and maintenance of auditory cortical asymmetries that does not depend on auditory language experience. Preservation of cortical volume in the deaf suggests plasticity in the input and output of auditory cortex that could include language-specific or more general-purpose information from other sensory modalities.     

Neural correlates of timbre change in harmonic sounds

Timbre is a major structuring force in music and one of the most important and ecologically relevant features of auditory events. We used sound stimuli selected on the basis of previous psychophysiological studies to investigate the neural correlates of timbre perception. Our results indicate that both the left and right hemispheres are involved in timbre processing, challenging the conventional notion that the elementary attributes of musical perception are predominantly lateralized to the right hemisphere. Significant timbre-related brain activation was found in well-defined regions of posterior Heschl's gyrus and superior temporal sulcus, extending into the circular insular sulcus. Although the extent of activation was not significantly different between left and right hemispheres, temporal lobe activations were significantly posterior in the left, compared to the right, hemisphere, suggesting a functional asymmetry in their respective contributions to timbre processing. The implications of our findings for music processing in particular and auditory processing in general are discussed.     

Abnormal association between reduced magnetic mismatch field to speech sounds and smaller left planum temporale volume in schizophrenia

Schizophrenia is associated with language-related dysfunction. A previous study [Schizophr. Res. 59 (2003c) 159] has shown that this abnormality is present at the level of automatic discrimination of change in speech sounds, as revealed by magnetoencephalographic recording of auditory mismatch field in response to across-category change in vowels. Here, we investigated the neuroanatomical substrate for this physiological abnormality. Thirteen patients with schizophrenia and 19 matched control subjects were examined using magnetoencephalography (MEG) and high-resolution magnetic resonance imaging (MRI) to evaluate both mismatch field strengths in response to change between vowel /a/ and /o/, and gray matter volumes of Heschl's gyrus (HG) and planum temporale (PT). The magnetic global field power of mismatch response to change in phonemes showed a bilateral reduction in patients with schizophrenia. The gray matter volume of left planum temporale, but not right planum temporale or bilateral Heschl's gyrus, was significantly smaller in patients with schizophrenia compared with that in control subjects. Furthermore, the phonetic mismatch strength in the left hemisphere was significantly correlated with left planum temporale gray matter volume in patients with schizophrenia only. These results suggest that structural abnormalities of the planum temporale may underlie the functional abnormalities of fundamental language-related processing in schizophrenia.     

Asymmetry in the human primary somatosensory cortex and handedness

Brain asymmetry is a phenomenon well known for handedness and language specialization and has also been studied in motor cortex. Less is known about hemispheric asymmetries in the somatosensory cortex. In the present study, we systematically investigated the representation of somatosensory function analyzing early subcortical and cortical somatosensory-evoked potentials (SEP) after electrical stimulation of the right and left median nerve. In 16 subjects, we compared thresholds, the peripheral neurogram at Erb point, and, using MRI-based EEG source analysis, the P14 brainstem component as well as N20 and P22, the earliest cortical responses from the primary sensorimotor cortex. Handedness was documented using the Edinburgh Inventory and a dichotic listening test was performed as a measure for language dominance. Whereas thresholds, Erb potential, and P14 were symmetrical, amplitudes of the cortical N20 showed significant hemispheric asymmetry. In the left hemisphere, the N20 amplitude was higher, its generator was located further medial, and it had a stronger dipole moment. There was no difference in dipole orientation. As a possible morphological correlate, the size of the left postcentral gyrus exceeded that of the right. The cortical P22 component showed a lower amplitude and a trend toward weaker dipole strength in the left hemisphere. Across subjects, there were no significant correlations between laterality indices of N20, the size of the postcentral gyrus, handedness, or ear advantage. These data show that asymmetry of median nerve SEP occurs at the cortical level, only. However, both functional and morphological cortical asymmetry of somatosensory representation appears to vary independently of motor and language functions.     

Bach speaks: a cortical "language-network" serves the processing of music

The aim of the present study was the investigation of neural correlates of music processing with fMRI. Chord sequences were presented to the participants, infrequently containing unexpected musical events. These events activated the areas of Broca and Wernicke, the superior temporal sulcus, Heschl's gyrus, both planum polare and planum temporale, as well as the anterior superior insular cortices. Some of these brain structures have previously been shown to be involved in music processing, but the cortical network comprising all these structures has up to now been thought to be domain-specific for language processing. To what extent this network might also be activated by the processing of non-linguistic information has remained unknown. The present fMRI-data reveal that the human brain employs this neuronal network also for the processing of musical information, suggesting that the cortical network known to support language processing is less domain-specific than previously believed.     

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.     

前阅读阶段儿童脑皮质结构变化与其语言发育的关系

目的 采用偏最小二乘法(PLS)观察前阅读阶段儿童皮质结构变化与其语言发育的关系。方法 于卡尔加里学龄前MRI研究成像数据库中选择来自39名健康发育儿童的117组脑结构T1WI,提取各ROI皮质指标,计算其偏侧化指数(AI)。记录儿童发育神经心理测评中语音处理和快速命名标准分数评价结果,以PLS回归分析观察语言能力与大脑皮质结构特征的关系。结果 语音标准处理分数与左侧环岛沟下段、左侧顶枕沟、左侧颞上沟、左侧中央下回和下沟、右侧额下回岛盖部皮质厚度均呈负相关,与前横侧副沟皮质厚度AI呈正相关。快速命名标准分数与胼胝体沟皮质厚度AI、大脑外侧眶沟和楔回高斯曲率AI均呈负相关,与大脑前横颞回、枕中沟和月状沟及嗅沟平均曲率AI均呈正相关。结论 语言中枢及附近脑区皮质结构改变可影响前阅读阶段儿童语言发育。     

Left thalamo-cortical network implicated in successful speech separation and identification

The separation of concurrent sounds is paramount to human communication in everyday settings. The primary auditory cortex and the planum temporale are thought to be essential for both the separation of physical sound sources into perceptual objects and the comparison of those representations with previously learned acoustic events. To examine the role of these areas in speech separation, we measured brain activity using event-related functional Magnetic Resonance Imaging (fMRI) while participants were asked to identify two phonetically different vowels presented simultaneously. The processing of brief speech sounds (200 ms in duration) activated the thalamus and superior temporal gyrus bilaterally, left anterior temporal lobe, and left inferior temporal gyrus. A comparison of fMRI signals between trials in which participants successfully identified both vowels as opposed to when only one of the two vowels was recognized revealed enhanced activity in left thalamus, Heschl's gyrus, superior temporal gyrus, and the planum temporale. Because participants successfully identified at least one of the two vowels on each trial, the difference in fMRI signal indexes the extra computational work needed to segregate and identify successfully the other concurrently presented vowel. The results support the view that auditory cortex in or near Heschl's gyrus as well as in the planum temporale are involved in sound segregation and reveal a link between left thalamo-cortical activation and the successful separation and identification of simultaneous speech sounds.     

The neural mechanism associated with the processing of onomatopoeic sounds

This event-related fMRI study was conducted to examine the blood-oxygen-level-dependent responses to the processing of auditory onomatopoeic sounds. We used a sound categorization task in which the participants heard four types of stimuli: onomatopoeic sounds, nouns (verbal), animal (nonverbal) sounds, and pure tone/noise (control). By discriminating between the categories of target sounds (birds/nonbirds), the nouns resulted in activations in the left anterior superior temporal gyrus (STG), whereas the animal sounds resulted in activations in the bilateral superior temporal sulcus (STS) and the left inferior frontal gyrus (IFG). In contrast, the onomatopoeias activated extensive brain regions, including the left anterior STG, the region from the bilateral STS to the middle temporal gyrus, and the bilateral IFG. The onomatopoeic sounds showed greater activation in the right middle STS than did the nouns and environmental sounds. These results indicate that onomatopoeic sounds are processed by extensive brain regions involved in the processing of both verbal and nonverbal sounds. Thus, we can posit that onomatopoeic sounds can serve as a bridge between nouns and animal sounds. This is the first evidence to demonstrate the way in which onomatopoeic sounds are processed in the human brain.     

Processing of sub-syllabic speech units in the posterior temporal lobe: an fMRI study

The objective of this study was to investigate phonological processing in the brain by using sub-syllabic speech units with rapidly changing frequency spectra. We used isolated stop consonants extracted from natural speech consonant-vowel (CV) syllables, which were digitized and presented through headphones in a functional magnetic resonance imaging (fMRI) paradigm. The stop consonants were contrasted with CV syllables. In order to control for general auditory activation, we used duration- and intensity-matched noise as a third stimulus category. The subjects were seventeen right-handed, healthy male volunteers. BOLD activation responses were acquired on a 1.5-T MR scanner. The auditory stimuli were presented through MR compatible headphones, using an fMRI paradigm with clustered volume acquisition and 12 s repetition time. The consonant vs. noise comparison resulted in unilateral left lateralized activation in the posterior part of the middle temporal gyrus and superior temporal sulcus (MTG/STS). The CV syllable vs. noise comparison resulted in bilateral activation in the same regions, with a leftward asymmetry. The reversed comparisons, i.e., noise vs. speech stimuli, resulted in right hemisphere activation in the supramarginal and superior temporal gyrus, as well as right prefrontal activation. Since the consonant stimuli are unlikely to have activated a semantic-lexical processing system, it seems reasonable to assume that the MTG/STS activation represents phonetic/phonological processing. This may involve the processing of both spectral and temporal features considered important for phonetic encoding.     

Controlling for individual differences in fMRI brain activation to tones, syllables, and words

Previous neuroimaging studies have consistently reported bilateral activation to speech stimuli in the superior temporal gyrus (STG) and have identified an anteroventral stream of speech processing along the superior temporal sulcus (STS). However, little attention has been devoted to the possible confound of individual differences in hemispheric dominance for speech. The present study was designed to test for speech-selective activation while controlling for inter-individual variance in auditory laterality, by using only subjects with at least 10% right ear advantage (REA) on the dichotic listening test. Eighteen right-handed, healthy male volunteers (median age 26) participated in the study. The stimuli were words, syllables, and sine wave tones (220-2600 Hz), presented in a block design. Comparing words > tones and syllables > tones yielded activation in the left posterior MTG and the lateral STG (upper bank of STS). In the right temporal lobe, the activation was located in the MTG/STS (lower bank). Comparing left and right temporal lobe cluster sizes from the words > tones and syllables > tones contrasts on single-subject level demonstrated a statistically significant left lateralization for speech sound processing in the STS/MTG area. The asymmetry analyses suggest that dichotic listening may be a suitable method for selecting a homogenous group of subjects with respect to left hemisphere language dominance.