Reading anomalous sentences: an event-related fMRI study of semantic processing

We report a random-effects analysis of an event-related fMRI study (n = 28) of cerebral activity during the reading of sentences that ended with a word that was either congruent or incongruent with the previous sentence context. Event-related potential studies have shown that this task elicits a late negativity peaking around 400 ms poststimulus (N400) that is larger for incongruent than for congruent sentence endings. A direct comparison of the activation for incongruent words versus that for congruent words revealed significantly greater activation for incongruent words than congruent words in bilateral inferior frontal and inferio-medial temporal cortex, left lateral frontal cortex, left posterior fusiform gyrus, bilateral motor cortex, and supplementary motor area. These results are consistent with data from intracranial electrical recording studies of the N400 electrical potential. The results are discussed as they relate to the localization of the cerebral sites underlying semantic processing in general and the localization of the scalp recorded N400 event-related potential in particular.     

Change detection in children with autism: an auditory event-related fMRI study

Autism involves impairments in communication and social interaction, as well as high levels of repetitive, stereotypic, and ritualistic behaviours, and extreme resistance to change. This latter dimension, whilst required for a diagnosis, has received less research attention. We hypothesise that this extreme resistance to change in autism is rooted in atypical processing of unexpected stimuli. We tested this using auditory event-related fMRI to determine regional brain activity associated with passive detection of infrequently occurring frequency-deviant and complex novel sounds in a no-task condition. Participants were twelve 10- to 15-year-old children with autism and a group of 12 age- and sex-matched healthy controls. During deviance detection, significant activation common to both groups was located in the superior temporal and inferior frontal gyri. During 'novelty detection', both groups showed activity in the superior temporal gyrus, the temporo-parietal junction, the superior and inferior frontal gyri, and the cingulate gyrus. Children with autism showed reduced activation of the left anterior cingulate cortex during both deviance and novelty detection. During novelty detection, children with autism also showed reduced activation in the bilateral temporo-parietal region and in the right inferior and middle frontal areas. This study confirms previous evidence from ERP studies of atypical brain function related to automatic change detection in autism. Abnormalities involved a cortical network known to have a role in attention switching and attentional resource distribution. These results throw light on the neurophysiological processes underlying autistic 'resistance to change'.     

Auditory processing of different types of pseudo-words: an event-related fMRI study

Imaging results on real word and pseudo-word processing have been heterogeneous, allowing only cautious claims about the neuroanatomical loci of lexico-semantic processing. In order to shed more light on this issue, we examined the impact of different structures of non-lexical stimuli on the outcome of comparisons between such items and matched real words. We anticipated that the degree to which a pseudo-word still resembles a particular real word template determines how word-like it is processed. To verify this idea, we tested different types of pseudo-words (either phonotactically legal and transparently or opaquely derived from real words or phonotactically illegal) in an event-related fMRI paradigm utilizing a lexical decision task. All types of pseudo-words elicited a stronger hemodynamic brain response than real words in the bilateral superior temporal gyri. Real words produced stronger brain activations than pseudo-words in the left posterior middle temporal and angular gyri, the rostral and caudal cingulate gyrus, the precuneus and the right inferior temporal gyrus. When contrasted to opaque pseudo-words transparent pseudo-words elicited a stronger brain response in a temporo-parietal region adjacent to the one observed for real words. Our results provide further support for the involvement of the left posterior middle temporal and angular gyri in lexical-semantic processing. The data also indicate that transparently derived pseudo-words are processed similarly to real words. In contrast, semantic operations are blocked when opaquely derived pseudo-words are processed.     

Combined event-related fMRI and intracerebral ERP study of an auditory oddball task

Event-related fMRI (efMRI) has been repeatedly used to seek the neural sources of endogenous event-related potentials (ERP). However, significant discrepancies exist between the efMRI data and the results of previously published intracranial ERP studies of oddball task. To evaluate the capacity of efMRI to define the sources of the P3 component of ERP within the human brain, both efMRI and intracerebral ERP recordings were performed in eight patients with intractable epilepsy (five males and three females) during their preoperative invasive video-EEG monitoring. An identical auditory oddball task with frequent and target stimuli was completed in two sessions. A total of 606 intracerebral sites were electrophysiologically investigated by means of depth electrodes. In accordance with the finding of multiple intracerebral generators of P3 potential, the target stimuli evoked MRI signal increase in multiple brain regions. However, regions with evident hemodynamic and electrophysiological responses overlapped only partially. P3 generators were always found within hemodynamic-active sites, if these sites were investigated by means of depth electrodes. On the other hand, unequivocal local sources of P3 potential were apparently also located outside the regions with a significant hemodynamic response (typically in mesiotemporal regions). Both methods should thus be viewed as mutually complementary in investigations of the spatial distribution of cortical and subcortical activation during oddball task.     

功能磁共振观察儿童文盲与非文盲语言加工相关脑区

目的 利用事件相关功能磁共振成像技术(fMRI)对儿童文盲和非文盲汉语加工相关脑区进行探讨.方法 分别对38名儿童(19名文盲和19名非文盲)进行汉字和简单图形判断的测试,同时采集其脑部的fMRI数据.使用基于Matlab7.0的SPM2软件进行数据分析,并生成脑功能区激活图像,对组内和组间数据分别进行单样本和双样本t检验分析.结果 儿童汉字加工激活了由多个脑区构成的神经网络,与成人基本一致,其中非文盲组和文盲组的差异性脑区位于左侧额中回(BA46/9),左侧角回(BA39),左侧梭状回(BA37).结论 左侧额中回(BA46/9)、左侧角回(BA39)及左侧梭状回(BA37)与汉语语义加工密切相关,读写能力在一定程度上改变了语言加工的神经网络联结模式.     

Frequency effects of Chinese character processing in the brain: an event-related fMRI study

Knowing how the brain processes Chinese characters of different frequencies of occurrence may shed light on the extent to which orthographic variations of different languages can influence reading processes in the brain. In the present study, event-related fMRI was used to investigate frequency effects on Chinese character processing. Reading low-frequency characters invoked higher activation in several brain regions including the left premotor/inferior frontal gyrus, supplementary motor area, left anterior insula, left posterior inferior temporal gyrus, left superior parietal cortex, and lingual cortex, while reading high-frequency characters resulted in higher activation in the left supramarginal/angular gyrus and left precuneus. The activation pattern of reading infrequently encountered characters reflects a more demanding processing procedure of retrieving, formulating, and coordinating the phonological output. Access to the lexical route may benefit the reading of high-frequency characters. By uncovering the differential brain responses in reading Chinese characters of different occurrence frequencies, not only has a substantial overlap between functional neuroanatomy of reading Chinese and alphabetical languages been demonstrated, but also features permitting the separation of language-specific content from universal mechanisms.     

Dissociating response conflict from numerical magnitude processing in the brain: an event-related fMRI study

Functional neuroimaging studies of numerical cognition have repeatedly associated activation of the intraparietal sulcus (IPS) with number processing. During number comparison, the IPS has been found to be modulated by the numerical distance. This has lead to the contention that the IPS houses the internal representation of numerical magnitude. However, this theory has been challenged by the argument that IPS activation may reflect domain-general response selection. In the present study, we used the numerical size congruity paradigm to further elucidate the role played by the IPS in number comparison. In an event-related, functional magnetic resonance imaging (fMRI) study, participants judged which of two number words was numerically larger. In addition to the numerical distance, physical stimulus size was varied such that physical size and numerical magnitude were either (a) congruent (e.g., numerically smaller number printed in smaller font) or (b) incongruent (e.g., numerically larger number printed in smaller font). This allowed for the study of both the main effects and the interaction of numerical distance and stimulus congruency. A main effect of numerical distance was found in bilateral regions of the IPS. However, these parietal areas were not significantly modulated by congruency or the interaction of distance and congruency. Instead, the main effect of congruency and an interaction of distance and congruency were observed in anterior cingulate and prefrontal cortices. These findings suggest some degree of independence between the processing of numerical distance and size congruity, lending support for the hypothesis that distance effects in IPS reflect the underlying representation of numerical magnitude.     

Feeling-of-knowing in episodic memory: an event-related fMRI study

An individual may fail to recall an item from memory but still feel that it would be recognized on a later test, a retrieval state termed the "feeling-of-knowing" (FOK). In this study we used event-related fMRI and the FOK to examine both encoding- and retrieval-related factors that are associated with different levels of recall performance: successful retrieval of a previously studied item, retrieval failure accompanied by the FOK, and retrieval failure without any FOK. The results revealed one predominant pattern of retrieval-related activation: an intermediate level of activation for FOK-less than that associated with successful recall and greater than that associated with unsuccessful recall (frontal and left parietal cortices). Two further patterns were also observed: greater activation for both successful recall and FOK than for unsuccessful recall (left midlateral prefrontal cortex) and greater activation for successful recall than for both FOK and unsuccessful recall (left MTL). Analysis of encoding trials conditional upon subsequent retrieval success revealed a pattern of activation that appeared to predict subsequent recall, but which further analysis indicated to be a better predictor of subsequent recognition. These results provide evidence that the phenomenology of graded recall is represented neurally in frontal and parietal cortices, but that activation at encoding may not precipitate the different levels of recall experience.     

The functional anatomy of inspection time: an event-related fMRI study

Twenty healthy young adults underwent functional magnetic resonance imaging (fMRI) of the brain while performing a visual inspection time task. Inspection time is a forced-choice, two-alternative visual backward-masking task in which the subject is briefly shown two parallel vertical lines of markedly different lengths and must decide which is longer. As stimulus duration decreases, performance declines to chance levels. Individual differences in inspection time correlate with higher cognitive functions. An event-related design was used. The hemodynamic (blood oxygenation level-dependent; BOLD) response was computed as both a function of the eight levels of stimulus duration, from 6 ms (where performance is almost at chance) to 150 ms (where performance is nearly perfect), and a function of the behavioral responses. Random effects analysis showed that the difficulty of the visual discrimination was related to bilateral activation in the inferior fronto-opercular cortex, superior/medial frontal gyrus, and anterior cingulate gyrus, and bilateral deactivation in the posterior cingulate gyrus and precuneus. Examination of the time courses of BOLD responses showed that activation was related specifically to the more difficult, briefer stimuli and that deactivation was found across most stimulus levels. Functional connectivity suggested the existence of two networks. One comprised the fronto-opercular area, intrasylvian area, medial frontal gyrus, and the anterior cingulate cortex (ACC), possibly associated with processing of visually degraded percepts. A posterior network of sensory-related and associative regions might subserve processing of a visual discrimination task that has high processing demands and combines several fundamental cognitive domains. fMRI can thus reveal information about the neural correlates of mental events which occur over very short durations.     

An event-related fMRI study of implicit phrase-level syntactic and semantic processing.

Most neuroimaging studies of language function to date use a block-subtraction paradigm in which images acquired during relatively long periods of target stimuli are compared to those acquired during a control period. These studies typically require an overt response on the part of the subject, usually some type of discrimination or grammatical judgment by button press, or silent word generation. Results from studies of syntactic and semantic processing have generally been compatible with the classical correlation to Broca's area and Wernicke's area, respectively. Recently, functional magnetic resonance imaging (fMRI) studies departing from the block-subtraction paradigm in favor of event-related fMRI paradigms have been reported. We have extended the use of this approach to examine implicit (i.e., without an explicit task on the part of the subject) syntactic and semantic processing at the phrasal level, using visually presented verb phrases. Left BA 44 is more strongly activated for the syntactic condition than the semantic condition. BA 45, 10, and 46 show laterality differences: mostly left-lateralized for the syntactic condition and right-lateralized for the semantic condition. We also find activations of the inferior parietal lobe, consistent with a visual oddball response reported previously, and the anterior cingulate gyrus (BA 32), implicated for attention and memory-related processes in numerous studies.     

Unilateral hemispheric lesions disrupt parallel processing within the contralateral intact hemisphere: an auditory fMRI study

Evidence from activation studies suggests that sound recognition and localization are processed in two distinct cortical networks that are each present in both hemispheres. Sound recognition and/or localization may, however, be disrupted by purely unilateral damage, suggesting that processing within one hemisphere may not be sufficient or may be disturbed by the contralateral lesion. Sound recognition and localization were investigated psychophysically and using fMRI in patients with unilateral right hemisphere lesions. Two patients had a combined deficit in sound recognition and sound localization, two a selective deficit in sound localization, one a selective deficit in sound recognition, and two normal performance in both tasks. The overall level of activation in the intact left hemisphere of the patients was smaller than in normal control subjects, irrespective of whether the patient's performance in the psychophysical tasks was impaired. Despite this overall decrease in activation strength, patients with normal performance still exhibited activation patterns similar to those of the control subjects in the recognition and localization tasks, indicating that the specialized brain networks subserving sound recognition and sound localization in normal subjects were also activated in the patients with normal performance, albeit to an altogether lesser degree. In patients with deficient performance, on the other hand, the activation patterns during the sound recognition and localization tasks were severely reduced, comprising fewer and partly atypical activation foci compared to the normal subjects. This indicates that impaired psychophysical performance correlates with a breakdown of parallel processing within specialized networks in the contralesional hemisphere.     

Amplitopicity of the human auditory cortex: an fMRI study

Whereas specialized frequency-encoding patterns in the human auditory cortex are generally accepted, termed tonotopicity, a similar principle of intensity encoding--amplitopicity--is debated controversially. This functional magnetic resonance imaging study describes the relationship of the activation volume and the spatial distribution of activated clusters under different sound pressure levels (SPL) across the temporal plane including the transverse temporal gyrus (TTG). Nine healthy subjects with no hearing deficiencies were investigated using an echo-planar imaging technique at 1.5 T. A boxcar stimulation paradigm was applied with a 5-Hz pulsed sine tone of 1000 Hz frequency at three SPLs of 70, 82, and 90 dB. Linear cross-correlation analysis (correlation coefficient > 0.3 corresponding to P < 0.08) of the functional data set revealed bilateral BOLD response within the auditory cortex of the nine subjects with moderate increase of activation volume for higher sound pressure levels. With increasing sound pressure a two-dimensional drift of cortical activation was observed (a) from the ventral to the dorsal edge and (b) from lateral to medial parts of TTG. This latero-medial drift therefore mimics the well-accepted principle of tonotopy for frequency-encoding neurons. This study demonstrates the existence of an amplitopic pattern of intensity-encoding neuronal clusters that in part resembles the tonotopic distribution of frequency-encoding neurons. This finding has to be integrated into the understanding of the auditory organization for the interpretation of higher auditory functions such as sound perception or speech.     

The effect of preceding context on inhibition: an event-related fMRI study

In this study we combined event-related fMRI with a parametric manipulation of the go nogo paradigm to examine the effect of preceding context on inhibitory processes. Nogo trials were preceded by either 1, 3, or 5 go trials and then compared to one another. Two distinct patterns of activation were associated with behavioral inhibition: First, the ventral prefrontal cortex, cingulate gyrus, and superior parietal regions showed a context effect with an increase in MR signal to nogo trials with increasing number of preceding go trials. Second, anterior regions in the supplementary and premotor cortex showed an increase in MR signal on the nogo condition after 5 preceding go trials, but not after only 1 or 3. A model using the BOLD response in our data was used to verify that the effect of context was not an artifact of the randomization scheme used in the design.     

Perirhinal cortex activity during visual object discrimination: an event-related fMRI study

Previous fMRI studies have demonstrated preferential involvement of the perirhinal cortex and hippocampus in tasks of object and spatial memory, respectively. Here we investigated whether similar activity would also be present when object and spatial discrimination was assessed in the absence of explicit declarative memory demands. On each trial in the scanner, participants were presented simultaneously with two arrays of objects and were asked to indicate whether both arrays were identical, differed with respect to the identity of one object or differed with respect to the spatial arrangement of the objects. It was found that the detection of an object identity change was associated with significant right perirhinal cortex activity. We suggest that this perirhinal activity indicates a role of this structure in processes beyond declarative memory, for example, short-term visual working memory or higher order perception. Significantly greater hippocampal activity was not, however, observed during the spatial arrangement condition, perhaps due to the relatively low spatial processing demands of this task.     

A fronto-parietal circuit for tactile object discrimination: an event-related fMRI study

Previous studies of somatosensory object discrimination have been focused on the primary and secondary sensorimotor cortices. However, we expected the prefrontal cortex to also become involved in sequential tactile discrimination on the basis of its role in working memory and stimulus discrimination as established in other domains. To investigate the contributions of the different cerebral structures to tactile discrimination of sequentially presented objects, we obtained event-related functional magnetic resonance images from seven healthy volunteers. Our results show that right hand object exploration involved left sensorimotor cortices, bilateral premotor, parietal and temporal cortex, putamen, thalamus, and cerebellum. Tactile exploration of parallelepipeds for subsequent object discrimination activated further areas in the dorsal and ventral portions of the premotor cortex, as well as parietal, midtemporal, and occipital areas of both cerebral hemispheres. Discriminating a parallelepiped from the preceding one involved a bilateral prefrontal-anterior cingulate-superior temporal-posterior parietal circuit. While the prefrontal cortex was active with right hemisphere dominance during discrimination, there was left hemispheric prefrontal activation during the delay period between object presentations. Delay related activity was further seen in the anterior intraparietal area and the fusiform gyrus. The results reveal a prominent role of the human prefrontal cortex for somatosensory object discrimination in correspondence with recent models on stimulus discrimination and working memory.     

Crinkling and crumpling: an auditory fMRI study of material properties

Knowledge of an object's material composition (i.e., what it is made of) alters how we interact with that object. Seeing the bright glint or hearing the metallic crinkle of a foil plate for example, confers information about that object before we have even touched it. Recent research indicates that the medial aspect of the ventral visual pathway is sensitive to the surface properties of objects. In the present functional magnetic resonance imaging (fMRI) study, we investigated whether the ventral pathway is also sensitive to material properties derived from sound alone. Relative to scrambled material sounds and non-verbal human vocalizations, audio recordings of materials being manipulated (i.e., crumpled) in someone's hands elicited greater BOLD activity in the right parahippocampal cortex of neurologically intact listeners, as well as a cortically blind participant. Additional left inferior parietal lobe activity was also observed in the neurologically intact group. Taken together, these results support a ventro-medial pathway that is specialized for processing the material properties of objects, and suggest that there are sub-regions within this pathway that subserve the processing of acoustically-derived information about material composition.     

Adults and children processing music: an fMRI study

The present study investigates the functional neuroanatomy of music perception with functional magnetic resonance imaging (fMRI). Three different subject groups were investigated to examine developmental aspects and effects of musical training: 10-year-old children with varying degrees of musical training, adults without formal musical training (nonmusicians), and adult musicians. Subjects made judgements on sequences that ended on chords that were music-syntactically either regular or irregular. In adults, irregular chords activated the inferior frontal gyrus, orbital frontolateral cortex, the anterior insula, ventrolateral premotor cortex, anterior and posterior areas of the superior temporal gyrus, the superior temporal sulcus, and the supramarginal gyrus. These structures presumably form different networks mediating cognitive aspects of music processing (such as processing of musical syntax and musical meaning, as well as auditory working memory), and possibly emotional aspects of music processing. In the right hemisphere, the activation pattern of children was similar to that of adults. In the left hemisphere, adults showed larger activations than children in prefrontal areas, in the supramarginal gyrus, and in temporal areas. In both adults and children, musical training was correlated with stronger activations in the frontal operculum and the anterior portion of the superior temporal gyrus.     

Self awareness and speech processing: an fMRI study

Language production and perception imply motor system recruitment. Therefore, language should obey the theory of shared motor representation between self and other, by means of mirror-like systems. These mirror-like systems (referring to single-unit recordings in animals) show the property to be recruited both when accomplishing and when perceiving a goal-directed action, whatever the sensory modality may be. This hypothesis supposes that a neural network for self-awareness is involved to distinguish speech production from speech listening. We used fMRI to test this assumption in 12 healthy subjects, who performed two different block-design experiments. The first experiment showed involvement of a lateral mirror-like network in speech listening, including ventral premotor cortex, superior temporal sulcus and the inferior parietal lobule (IPL). The activity of this mirror-like network is associated with the perception of an intelligible speech. The second experiment looked at a self-awareness network. It showed involvement of a medial resting-state network, including the medial parietal and medial prefrontal cortices, during the 'self-generated voice' condition, as opposed to passive speech listening. Our results support the fact that deactivation of this medial network, in association with modulation of the activity of the IPL (part of the mirror-like network previously described), is linked to self-awareness in speech processing. Overall, these results support the idea that self-awareness is present when distinguishing between speech production and speech listening situations, and may depend on these two different parieto-frontal networks.     

Volition to action--an event-related fMRI study

Current concepts of the anterior cingulate cortex (ACC) increasingly emphasize its role as an interface between limbic and neocortical functions. It has been pointed out that ACC activation reflects the intentional amount of effort (volition) that a subject uses in a task. In previous electrophysiological source localization investigations during a choice reaction task, we described a strong early activation in the ACC region approximately 120-150 ms after stimulus presentation. The degree of midline ACC activation correlated negatively with reaction time. This observation together with the finding that ACC activation precedes information processing in cortical association areas provided preliminary support to the notion that the extent of ACC activation is related to a subject's task engagement. However, due to the inverse problem and the relatively low spatial resolution of the electrophysiological measurements, we were not able to make inferences about the validity and the exact localization of the observed midline activation maximum. We addressed this question and performed an event-related fMRI study in six healthy volunteers during a visual choice reaction task. Two checkerboard stimuli were presented either in the left or right visual hemifield in randomized order and with an interstimulus interval requiring an appropriate motor response (left-right button press). A bilateral BOLD maximum was observed in the region of the supplementary motor area confluent with the neighboring motor area of the dorsal ACC. The degree of ACC activation correlated significantly with reaction time. These results are in line with our previous electrophysiological findings and provide further evidence that early ACC activation during a choice reaction task reflects the intentional effort of a subject to carry out a task.     

Semantic ambiguity processing in sentence context: Evidence from event-related fMRI

Lexical semantic ambiguity is the phenomenon when a word has multiple meanings (e.g. 'bank'). The aim of this event-related functional MRI study was to identify those brain areas, which are involved in contextually driven ambiguity resolution. Ambiguous words were selected which have a most frequent, dominant, and less frequent, subordinate meaning. These words were presented in two types of sentences: (1) a sentence congruent with the dominant interpretation and (2) a sentence congruent with the subordinate interpretation. Sentences without ambiguous words served as a control condition. The ambiguous words always occurred early in the sentences and were biased towards one particular meaning by the final word(s) of the sentence; the event at the end of the sentences was modeled. The results indicate that a bilaterally distributed network supports semantic ambiguity comprehension: left (BA 45/44) and right (BA 47) inferior frontal gyri and left (BA 20/37) and right inferior/middle temporal gyri (BA 20). The pattern of activation is most consistent with a scenario in which initially a frequency-based probabilistic choice is made between the alternative meanings, and the meaning is updated when this interpretation does not fit into the final disambiguating context. The neural pattern is consistent with the results of other neuroimaging experiments which manipulated various aspects of integrative and context processing task demands. The presence of a bilateral network is also in line with the lesion and divided visual field literature, but contrary to earlier claims, the two hemispheres appear to play similar roles during semantic ambiguity resolution.