Rhythmic brain activities related to singing in humans

To investigate the motor control related to sound production, we studied cortical rhythmic changes during continuous vocalization such as singing. Magnetoencephalographic (MEG) responses were recorded while subjects spoke in the usual way (speaking), sang (singing), hummed (humming) and imagined (imagining) a popular song. The power of alpha (8-15 Hz), beta (15-30 Hz) and low-gamma (30-60 Hz) frequency bands was changed during and after vocalization (singing, speaking and humming). In the alpha band, the oscillatory changes for singing were most pronounced in the right premotor, bilateral sensorimotor, right secondary somatosensory and bilateral superior parietal areas. The beta oscillation for the singing was also confirmed in the premotor, primary and secondary sensorimotor and superior parietal areas in the left and right hemispheres where were partly activated even for imagined a song (imaging). These regions have been traditionally described as vocalization-related sites. The cortical rhythmic changes were distinct in the singing condition compared with the other vocalizing conditions (speaking and humming) and thus we considered that more concentrated control of the vocal tract, diaphragm and abdominal muscles is responsible. Furthermore, characteristic oscillation in the high-gamma (60-200 Hz) frequency band was found in Broca's area only in the imaging condition and might occur singing rehearsal and storage process in Broca's area.     

Developmental changes in the neural correlates of semantic processing

Functional magnetic resonance imaging (fMRI) was used to explore the neural correlates of semantic judgments in the auditory modality in a group of 9- to 15-year-old children. Subjects were required to indicate if word pairs were related in meaning. Consistent with previous findings in adults, children showed activation in bilateral superior temporal gyri (BA 22) for recognizing spoken words as well as activations in bilateral inferior frontal gyri (BAs 47, 45) and left middle temporal gyrus (BA 21) for semantic processing. The neural substrates of semantic association and age differences were also investigated. Words with strong semantic association elicited significantly greater activation in the left inferior parietal lobule (BA 40), whereas words with weak semantic association elicited activation in left inferior frontal gyrus (BAs 47/45). Correlations with age were observed in the left middle temporal gyrus (BA 21) and the right inferior frontal gyrus (BA 47). The pattern of results for semantic association implies that the left inferior parietal lobule effectively integrates highly related semantic features and the left inferior frontal gyrus becomes more active for words that require a greater search for semantic associations. The developmental results suggest that older children recruit the right inferior frontal gyrus as they conduct a broader semantic search and the left middle temporal gyrus to provide more efficient access to semantic representations.     

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.     

Experience-dependent neural substrates involved in vocal pitch regulation during singing

Proper singing requires the integration of auditory feedback mechanisms with the vocal motor system, such that vocal pitch can be precisely controlled. To determine the neural substrates involved in audio-vocal integration, non-musicians and experienced singers underwent fMRI scanning while they sang a single tone with either unaltered (“simple”) or pitch-shifted auditory feedback; in pitch-shifted trials, subjects were instructed either to ignore or compensate for the shifted feedback. We hypothesized that the anterior cingulate cortex (ACC), superior temporal gyrus (STG), and anterior insula may be involved in audio-vocal integration due to their functional roles during singing and their anatomical connectivity. Although singers were more accurate than non-musicians in simple singing, both groups recruited similar functional networks. Singers ignored the shifted feedback better than non-musicians, and both groups also displayed different patterns of neural activity for this task: singers recruited bilateral auditory areas and left putamen, while non-musicians recruited the left supramarginal gyrus and primary motor cortex. While there were no significant group differences in performing the compensate task, singers displayed enhanced activity in the ACC, superior temporal sulcus, and putamen, whereas non-musicians exhibited increased activity in the dorsal premotor cortex, a region involved with sensorimotor interactions. We propose two neural substrates for audio-vocal integration: the dorsal premotor cortex may act as a basic interface, but with vocal training and practice, the ACC, auditory cortices, and putamen may be increasingly recruited as people learn to monitor their auditory feedback and adjust their vocal output accordingly.     

中英文语言活动区功能磁共振成像研究

目的初步探讨母语为中文者,说中文(L1)及英文(L2)时激活的语言相关功能区并比较其异同。方法对22例正常志愿者———“晚双语(13～15岁开始学英语)中国人,于说中文及英文两种状态下分别进行血氧水平依赖性增强效应磁共振脑功能成像检查,研究哪些语言相关脑功能区被激活,并比较两种状态下的脑激活区。结果说中文及英文时分别激活了不完全相同的脑区。说中文时激活的主要脑区包括:两侧运动区,左右侧额下回,左右侧颞上回,左侧岛叶及左右侧小脑半球。说英文时激活的脑区包括:左侧额下回,左侧颞上回,左侧岛叶及左右侧小脑半球。比较显示中文任务可激活右侧颞上回,而英文任务未激活这些区域。结论中英文“晚双语”者,两种语言激活的脑区不完全相同。说中文时激活的脑区要比说英文时较广泛。这主要在右侧大脑半球的右额下回及右颞上回。     

Towards a fronto-temporal neural network for the decoding of angry vocal expressions

Vocal expressions commonly elicit activity in superior temporal and inferior frontal cortices, indicating a distributed network to decode vocally expressed emotions. We examined the involvement of this fronto-temporal network for the decoding of angry voices during attention towards (explicit attention) or away from emotional cues in voices (implicit attention) based on a reanalysis of previous data (Frühholz, S., Ceravolo, L., Grandjean, D., 2012. Cerebral Cortex 22, 1107-1117). The general network revealed high interconnectivity of bilateral inferior frontal gyrus (IFG) to different bilateral voice-sensitive regions in mid and posterior superior temporal gyri. Right superior temporal gyrus (STG) regions showed connectivity to the left primary auditory cortex and secondary auditory cortex (AC) as well as to high-level auditory regions. This general network revealed differences in connectivity depending on the attentional focus. Explicit attention to angry voices revealed a specific right-left STG network connecting higher-level AC. During attention to a nonemotional vocal feature we also found a left-right STG network implicitly elicited by angry voices that also included low-level left AC. Furthermore, only during this implicit processing there was widespread interconnectivity between bilateral IFG and bilateral STG. This indicates that while implicit attention to angry voices recruits extended bilateral STG and IFG networks for the sensory and evaluative decoding of voices, explicit attention to angry voices solely involves a network of bilateral STG regions probably for the integrative recognition of emotional cues from voices.     

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.     

Overlapping neural regions for processing rapid temporal cues in speech and nonspeech signals

Speech perception involves recovering the phonetic form of speech from a dynamic auditory signal containing both time-varying and steady-state cues. We examined the roles of inferior frontal and superior temporal cortex in processing these aspects of auditory speech and nonspeech signals. Event-related functional magnetic resonance imaging was used to record activation in superior temporal gyrus (STG) and inferior frontal gyrus (IFG) while participants discriminated pairs of either speech syllables or nonspeech tones. Speech stimuli differed in either the consonant or the vowel portion of the syllable, whereas the nonspeech signals consisted of sinewave tones differing along either a dynamic or a spectral dimension. Analyses failed to identify regions of activation that clearly contrasted the speech and nonspeech conditions. However, we did identify regions in the posterior portion of left and right STG and left IFG yielding greater activation for both speech and nonspeech conditions that involved rapid temporal discrimination, compared to speech and nonspeech conditions involving spectral discrimination. The results suggest that, when semantic and lexical factors are adequately ruled out, there is significant overlap in the brain regions involved in processing the rapid temporal characteristics of both speech and nonspeech signals.     

Development of brain networks involved in spoken word processing of Mandarin Chinese

Developmental differences in phonological and orthographic processing of Chinese spoken words were examined in 9-year-olds, 11-year-olds and adults using functional magnetic resonance imaging (fMRI). Rhyming and spelling judgments were made to two-character words presented sequentially in the auditory modality. Developmental comparisons between adults and both groups of children combined showed that age-related changes in activation in visuo-orthographic regions depended on a task. There were developmental increases in the left inferior temporal gyrus and the right inferior occipital gyrus in the spelling task, suggesting more extensive visuo-orthographic processing in a task that required access to these representations. Conversely, there were developmental decreases in activation in the left fusiform gyrus and left middle occipital gyrus in the rhyming task, suggesting that the development of reading is marked by reduced involvement of orthography in a spoken language task that does not require access to these orthographic representations. Developmental decreases may arise from the existence of extensive homophony (auditory words that have multiple spellings) in Chinese. In addition, we found that 11-year-olds and adults showed similar activation in the left superior temporal gyrus across tasks, with both groups showing greater activation than 9-year-olds. This pattern suggests early development of perceptual representations of phonology. In contrast, 11-year-olds and 9-year-olds showed similar activation in the left inferior frontal gyrus across tasks, with both groups showing weaker activation than adults. This pattern suggests late development of controlled retrieval and selection of lexical representations. Altogether, this study suggests differential effects of character acquisition on development of components of the language network in Chinese as compared to previous reports on alphabetic languages.     

Developmental changes in brain regions involved in phonological and orthographic processing during spoken language processing

Developmental differences in brain activation of 9- to 15-year-old children were examined during an auditory rhyme decision task to spoken words using functional magnetic resonance imaging (fMRI). As a group, children showed activation in the left superior/middle temporal gyri (BA 22, 21), right middle temporal gyrus (BA 21), dorsal (BA 45, pars opercularis) and ventral (BA 46, pars triangularis) aspects of the left inferior frontal gyrus, and left fusiform gyrus (BA 37). There was a developmental increase in activation in the left middle temporal gyrus (BA 22) across all lexical conditions, suggesting that automatic semantic processing increases with age regardless of task demands. Activation in the left dorsal inferior frontal gyrus also showed developmental increases for the conflicting (e.g. PINT-MINT) compared to the non-conflicting (e.g. PRESS-LIST) non-rhyming conditions, indicating that this area becomes increasingly involved in strategic phonological processing in the face of conflicting orthographic and phonological representations. Left inferior temporal/fusiform gyrus (BA 37) activation was also greater for the conflicting (e.g. PINT-MINT) condition, and a developmental increase was found in the positive relationship between individuals' reaction time and activation in the left lingual/fusiform gyrus (BA 18) in this condition, indicating an age-related increase in the association between longer reaction times and greater visual-orthographic processing in this conflicting condition. These results suggest that orthographic processing is automatically engaged by children in a task that does not require access to orthographic information for correct performance, especially when orthographic and phonological representations conflict, and especially for longer response latencies in older children.     

Processing of vocalizations in humans and monkeys: a comparative fMRI study

Humans and many other animals use acoustical signals to mediate social interactions with conspecifics. The evolution of sound-based communication is still poorly understood and its neural correlates have only recently begun to be investigated. In the present study, we applied functional MRI to humans and macaque monkeys listening to identical stimuli in order to compare the cortical networks involved in the processing of vocalizations. At the first stages of auditory processing, both species showed similar fMRI activity maps within and around the lateral sulcus (the Sylvian fissure in humans). Monkeys showed remarkably similar responses to monkey calls and to human vocal sounds (speech or otherwise), mainly in the lateral sulcus and the adjacent superior temporal gyrus (STG). In contrast, a preference for human vocalizations and especially for speech was observed in the human STG and superior temporal sulcus (STS). The STS and Broca's region were especially responsive to intelligible utterances. The evolution of the language faculty in humans appears to have recruited most of the STS. It may be that in monkeys, a much simpler repertoire of vocalizations requires less involvement of this temporal territory.     

Inhibit yourself and understand the other: neural basis of distinct processes underlying Theory of Mind

Taking the perspective of somebody else (Theory of Mind; ToM) is an essential human ability depending on a large cerebral network comprising prefrontal and temporo-parietal regions. Recently, ToM was suggested to consist of two processes: (1) self-perspective inhibition and (2) belief reasoning. Moreover, it has been hypothesized that self-perspective inhibition may build upon basic motor response inhibition. This study tested both hypotheses for the first time using functional Magnetic Resonance Imaging (fMRI), through administering both a ToM and a stop-signal paradigm in the same subjects. Both self-perspective and motor response inhibition yielded bilateral inferior frontal gyrus (IFG) activation, suggesting a common inhibitory mechanism, while belief reasoning was mediated by the superior temporal gyrus (STG) and temporo-parietal junction (TPJ). Thus, we provide neurobiological evidence for a subdivision of ToM into self-perspective inhibition and belief reasoning. Furthermore, evidence for partially shared neural mechanisms for inhibition in complex social situations and basic motor response inhibition was found.     

Developmental changes in activation and effective connectivity in phonological processing

The current study examined developmental changes in activation and effective connectivity among brain regions during a phonological processing task, using fMRI. Participants, ages 9-15, were scanned while performing rhyming judgments on pairs of visually presented words. The orthographic and phonological similarity between words in the pair was independently manipulated, so that rhyming judgment could not be based on orthographic similarity. Our results show a developmental increase in activation in the dorsal part of left inferior frontal gyrus (IFG), accompanied by a decrease in the dorsal part of left superior temporal gyrus (STG). The coupling of dorsal IFG with other selected brain regions involved in the phonological decision increased with age, while the coupling of STG decreased with age. These results suggest that during development there is a shift from reliance on sensory auditory representations to reliance on phonological segmentation and covert articulation for performing rhyming judgment on visually presented words. In addition, we found a developmental increase in activation in left posterior parietal cortex that was not accompanied by a change in its connectivity with the other regions. These results suggest that maturational changes within a cortical region are not necessarily accompanied by an increase in its interactions with other regions and its contribution to the task. Our results are consistent with the idea that there is reduced reliance on primary sensory processes as task-relevant processes mature and become more efficient during development.     

Effects of feature-selective attention on auditory pattern and location processing

Recent neuroimaging studies have suggested that spatial versus nonspatial changes in acoustic stimulation are processed along separate cortical pathways. However, it has remained unclear in how far change-related responses are modulated by selective attention. Thus, we aimed at testing effects of feature-selective attention on the cortical representation of pattern and location of complex natural sounds using human functional magnetic resonance imaging (fMRI) adaptation. We consecutively presented the following pairs of animal vocalizations: (a) two identical animal vocalizations, (b) same animal vocalizations at different locations, (c) different animal vocalizations at the same location, and (d) different animal vocalizations at different locations. Subjects underwent this stimulation under two different task conditions requiring either to match sound identity or location. We observed significant fMRI adaptation effects within the bilateral superior temporal sulcus (STS), planum temporale (PT) and right anterior insula for location changes. For pattern changes, we found adaptation effects within the bilateral superior temporal lobe, in particular along the superior temporal gyrus (STG), PT and posterior STS, the bilateral anterior insula and inferior frontal areas. While the adaptation effects within the pattern-selective temporal lobe areas were robust to task requirements, adaptation within the more posterior location-selective areas was modulated by feature-specific attention. In contrast, inferior frontal cortex and anterior insular exhibited adaptation effects mainly during the location matching task. Given that the location matching task was significantly more difficult than the pattern matching, our data suggest that frontal and insular regions were modulated by task difficulty rather than feature-specific attention.     

阅读半规则汉字时的脑激活模式:声调的作用

目的引入声调因素,运用功能磁共振成像技术进一步研究汉字规则性效应的脑激活模式.方法13名被试在磁共振扫描过程中执行视觉方式呈现的汉字出声阅读任务,根据形声字声旁与整字间读音及声调相同与否,将刺激材料细分为同音同调字、同音异调字、异音同调字和异音异调字,每类20字.结果四种条件均激活双侧额中回、额下回、前运动区及辅助运动区、左侧顶下小叶、双侧颞上回、双侧岛叶和右侧小脑,同时还各自激活了梭状回、颞中回、左内侧额叶等区域.另外,本实验还观察到这四种条件在右侧颞上回、双侧额下回等区域激活模式的特征性变化.结论存在汉字加工的半规则效应,声调的作用不应忽略.     

The brain basis of syntactic processes: functional imaging and lesion studies

Language comprehension can be subdivided into three processing steps: initial structure building, semantic integration, and late syntactic integration. The two syntactic processing phases are correlated with two distinct components in the event-related brain potential, namely an early left anterior negativity (ELAN) and a late centroparietal positivity (P600). Moreover, ERP findings from healthy adults suggest that early structure-building processes as reflected by the ELAN are independent of semantic processes. fMRI results have revealed that semantic and syntactic processes are supported by separable temporofrontal networks, with the syntactic processes involving the left superior temporal gyrus (STG), the left frontal operculum, and the basal ganglia (BG) in particular. MEG data from healthy adults have indicated that the left anterior temporal region and the left inferior frontal region subserve the early structure building processes. ERP data from patients with lesions in the left anterior temporal region and from patients with lesions in the left inferior frontal gyrus support this view, as these patients do not demonstrate an ELAN, although they do demonstrate a P600. Further results from patients with BG dysfunction suggest that parts of this subcortical structure are involved in late syntactic integrational processes. The data from the different experiments lead to the notion of separable brain systems responsible for early and late syntactic processes, with the former being subserved by the inferior frontal gyrus and the anterior STG and the latter being supported by the BG and more posterior portions of the STG.     

A systematic investigation of the functional neuroanatomy of auditory and visual phonological processing

Neuroimaging studies of auditory and visual phonological processing have revealed activation of the left inferior and middle frontal gyri. However, because of task differences in these studies (e.g., consonant discrimination versus rhyming), the extent to which this frontal activity is due to modality-specific linguistic processes or to more general task demands involved in the comparison and storage of stimuli remains unclear. An fMRI experiment investigated the functional neuroanatomical basis of phonological processing in discrimination and rhyming tasks across auditory and visual modalities. Participants made either "same/different" judgments on the final consonant or rhyme judgments on auditorily or visually presented pairs of words and pseudowords. Control tasks included "same/different" judgments on pairs of single tones or false fonts and on the final member in pairs of sequences of tones or false fonts. Although some regions produced expected modality-specific activation (i.e., left superior temporal gyrus in auditory tasks, and right lingual gyrus in visual tasks), several regions were active across modalities and tasks, including posterior inferior frontal gyrus (BA 44). Greater articulatory recoding demands for processing of pseudowords resulted in increased activation for pseudowords relative to other conditions in this frontal region. Task-specific frontal activation was observed for auditory pseudoword final consonant discrimination, likely due to increased working memory demands of selection (ventrolateral prefrontal cortex) and monitoring (mid-dorsolateral prefrontal cortex). Thus, the current study provides a systematic comparison of phonological tasks across modalities, with patterns of activation corresponding to the cognitive demands of performing phonological judgments on spoken and written stimuli.     

Exploring the visual world: the neural substrate of spatial orienting

Inspecting the visual environment, humans typically direct their attention across space by means of voluntary saccadic eye movements. Neuroimaging studies in healthy subjects have identified the superior parietal cortex and intraparietal sulcus as important structures involved in visual search. However, in apparent contrast, spatial disturbance of free exploration typically is observed after damage of brain structures located far more ventrally. Lesion studies in such patients disclosed the inferior parietal lobule (IPL) and temporo-parietal junction (TPJ), the superior temporal gyrus (STG) and insula, as well as the inferior frontal gyrus (IFG) of the right hemisphere. Here we used functional magnetic resonance imaging to investigate the involvement of these areas in active visual exploration in the intact brain. We conducted a region of interest analysis comparing free visual exploration of a dense stimulus array with the execution of stepwise horizontal and vertical saccades. The comparison of BOLD responses revealed significant signal increases during exploration in TPJ, STG, and IFG. This result calls for a reappraisal of the previous thinking on the function of these areas in visual search processes. In agreement with lesion studies, the data suggest that these areas are part of the network involved in human spatial orienting and exploration. The IPL dorsally of TPJ seem to be of minor importance for free visual exploration as these areas appear to be equally involved in the execution of spatially predetermined saccades.     

High frequency rTMS modulation of the sensorimotor networks: behavioral changes and fMRI correlates

Repetitive transcranial magnetic stimulation (rTMS) to the primary motor cortex (M1) may induce functional modulation of motor performance and sensory perception. To address the underlying neurophysiological modulation following 10 Hz rTMS applied over M1, we examined cortical activation using 3T functional magnetic resonance imaging (fMRI), as well as the associated motor and sensory behavioral changes. The motor performance measure involved a sequential finger motor task that was also used as an activation task during fMRI. For sensory assessment, current perception threshold was measured before and after rTMS outside the MR scanner, and noxious mechanical stimulation was used as an activation task during fMRI. We found that significant activation in the bilateral basal ganglia, left superior frontal gyrus, bilateral pre-SMA, right medial temporal lobe, right inferior parietal lobe, and right cerebellar hemisphere correlated with enhanced motor performance in subjects that received real rTMS compared with sham-stimulated controls. Conversely, significant deactivation in the right superior and middle frontal gyri, bilateral postcentral and bilateral cingulate gyri, left SMA, right insula, right basal ganglia, and right cerebellar hemisphere were associated with an increase in the sensory threshold. Our findings reveal that rTMS induced rapid changes in the sensorimotor networks associated with sensory perception and motor performance and demonstrate the complexity of such intervention.