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Background

We used fMRI to examine functional brain abnormalities of German-speaking dyslexics who suffer from slow effortful reading but not from a reading accuracy problem. Similar to acquired cases of letter-by-letter reading, the developmental cases exhibited an abnormal strong effect of length (i.e., number of letters) on response time for words and pseudowords.

Results

Corresponding to lesions of left occipito-temporal (OT) regions in acquired cases, we found a dysfunction of this region in our developmental cases who failed to exhibit responsiveness of left OT regions to the length of words and pseudowords. This abnormality in the left OT cortex was accompanied by absent responsiveness to increased sublexical reading demands in phonological inferior frontal gyrus (IFG) regions. Interestingly, there was no abnormality in the left superior temporal cortex which—corresponding to the onological deficit explanation—is considered to be the prime locus of the reading difficulties of developmental dyslexia cases.

Conclusions

The present functional imaging results suggest that developmental dyslexia similar to acquired letter-by-letter reading is due to a primary dysfunction of left OT regions.

We report three behavioral experiments on the spatial characteristics evoking illusory face and letter detection. False detections made to pure noise images were analyzed using a modified reverse correlation method in which hundreds of observers rated a modest number of noise images (480) during a single session. This method was originally developed for brain imaging research, and has been used in a number of fMRI publications, but this is the first report of the behavioral classification images. In Experiment 1 illusory face detection occurred in response to scattered dark patches throughout the images, with a bias to the left visual field. This occurred despite the use of a fixation cross and expectations that faces would be centered. In contrast, illusory letter detection (Experiment 2) occurred in response to centrally positioned dark patches. Experiment 3 included an oval in all displays to spatially constrain illusory face detection. With the addition of this oval the classification image revealed an eyes/nose/mouth pattern. These results suggest that face detection is triggered by a minimal face-like pattern even when these features are not centered in visual focus.

This fMRI study contrasted case-deviant and letter-deviant forms with familiar forms of the same phonological words (e.g., TaXi and Taksi vs. Taxi) and found, that both types of deviance led to increased activation in a left occipitotemporal region corresponding to the Visual Word Form Area. Case-deviant items, in addition, led to increased activation in a right occipitotemporal region and in a left occipital and a left posterior occipitotemporal region, possibly reflecting the increased demands on letter form coding. For letter-deviant items, in addition to the increased left occipitotemporal activation, a main finding was increased activation primarily in extended left frontal regions, possibly reflecting sublexically mediated access to word phonology. These findings are consistent with general features of cognitive dual-route models of visual word processing. Furthermore, they add support to the main feature of Dehaene et al.’s (2005) neural model of early stages of visual word processing . However, the increased activation found for case-deviant items in the VWFA cannot be immediately reconciled with the assumption of completely abstract case-independent orthographic word codes in the VWFA.

Previous studies have shown that developmental changes in the structure and function of prefrontal regions can continue throughout childhood and adolescence. Our recent results suggested a role for the left inferior frontal cortex in modulating task-dependent shifts in effective connectivity when adults focus on orthographic versus phonological aspects of presented words. Specifically, the top-down influence of the inferior frontal cortex determined whether incoming word-form information from the fusiform gyrus would have a greater impact on the parietal areas involved in orthographic processing or temporal areas involved in phonological processing. In the current study, we find that children displayed an identical pattern of task-dependent functional activations within this network. In comparison to adults, however, children had significantly weaker top-down modulatory influences emanating from the inferior frontal area. Adult language processing may thus involve greater top-down cognitive control compared to children, resulting in less interference from task-irrelevant information.

Evidence suggests that the neural system associated with face processing is a distributed cortical network containing both bottom-up and top-down mechanisms. While bottom-up face processing has been the focus of many studies, the neural areas involved in the top-down face processing have not been extensively investigated due to difficulty in isolating top-down influences from the bottom-up response engendered by presentation of a face. In the present study, we used a novel experimental method to induce illusory face detection. This method allowed for directly examining the neural systems involved in top-down face processing while minimizing the influence of bottom-up perceptual input. A distributed cortical network of top-down face processing was identified by analyzing the functional connectivity patterns of the right fusiform face area (FFA). This distributed cortical network model for face processing includes both “core” and “extended” face processing areas. It also includes left anterior cingulate cortex (ACC), bilateral orbitofrontal cortex (OFC), left dorsolateral prefrontal cortex (DLPFC), left premotor cortex, and left inferior parietal cortex. These findings suggest that top-down face processing contains not only regions for analyzing the visual appearance of faces, but also those involved in processing low spatial frequency (LSF) information, decision making, and working memory.

Individuals learn to read by gradually recognizing repeated letter combinations. However, it is unclear how or when neural mechanisms associated with repetition of basic stimuli (i.e., strings of letters) shift to involvement of higher-order language networks. The present study investigated this question by repeatedly presenting unfamiliar letter strings in a one-back matching task during an hour-long period. Activation patterns indicated that only brain areas associated with visual processing were activated during the early period, but additional regions that are usually associated with semantic and phonological processing in inferior frontal gyrus were recruited after stimuli became more familiar. Changes in activation were also observed in bilateral superior temporal cortex, also suggestive of a shift toward a more language-based processing strategy. Connectivity analyses reveal two distinct networks that correspond to phonological and visual processing, which may reflect the indirect and direct routes of reading. The phonological route maintained a similar degree of connectivity throughout the experiment, whereas visual areas increased connectivity with language areas as stimuli became more familiar, suggesting early recruitment of the direct route. This study provides insight about plasticity of the brain as individuals become familiar with unfamiliar combinations of letters (i.e., words in a new language, new acronyms) and has implications for engaging these linguistic networks during development of language remediation therapies.

Selective attention to speech versus nonspeech signals in complex auditory input could produce top-down modulation of cortical regions previously linked to perception of spoken, and even visual, words. To isolate such top-down attentional effects, we contrasted 2 equally challenging active listening tasks, performed on the same complex auditory stimuli (words overlaid with a series of 3 tones). Instructions required selectively attending to either the speech signals (in service of rhyme judgment) or the melodic signals (tone-triplet matching). Selective attention to speech, relative to attention to melody, was associated with blood oxygenation level–dependent (BOLD) increases during functional magnetic resonance imaging (fMRI) in left inferior frontal gyrus, temporal regions, and the visual word form area (VWFA). Further investigation of the activity in visual regions revealed overall deactivation relative to baseline rest for both attention conditions. Topographic analysis demonstrated that while attending to melody drove deactivation equivalently across all fusiform regions of interest examined, attending to speech produced a regionally specific modulation: deactivation of all fusiform regions, except the VWFA. Results indicate that selective attention to speech can topographically tune extrastriate cortex, leading to increased activity in VWFA relative to surrounding regions, in line with the well-established connectivity between areas related to spoken and visual word perception in skilled readers.

Imaging studies show that in normal language correlated activity between anterior and posterior brain regions increases as the linguistic and semantic content (i.e., from false fonts, letter strings, pseudo words, to words) of stimuli increase. In schizophrenia however, disrupted functional connectivity between frontal and posterior brain regions has been frequently reported and these disruptions may change the nature of language organization. We characterized basic linguistic operations in word and letter string processing in a region-of-interest network using structural equation modeling (SEM). Healthy volunteers and volunteers with schizophrenia performed an fMRI one-back matching task with real words and consonant letter strings. We hypothesized that left hemisphere network dysfunction in schizophrenia would be present during processes dealing with linguistic/semantic content. The modeling results suggest aberrant left hemisphere function in schizophrenia, even in tasks requiring minimal access to language. Alternative mechanisms included increases in right hemisphere involvement and increased top-down influence from frontal to posterior regions.

fMRI was used to examine lexical processing in native adult Chinese speakers. A 2 task (semantics and phonology) × 2 modality (visual and auditory) within-subject design was adopted. The semantic task involved a meaning association judgment and the phonological task involved a rhyming judgment to two sequentially presented words. The overall effect across tasks and modalities was used to identify seven ROIs, including the left fusiform gyrus (FG), the left superior temporal gyrus (STG), the left ventral inferior frontal gyrus (VIFG), the left middle temporal gyrus (MTG), the left dorsal inferior frontal gyrus (DIFG), the left inferior parietal lobule (IPL), and the left middle frontal gyrus (MFG). ROI analyses revealed two modality-specific areas, FG for visual and STG for auditory, and three task-specific areas, IPL and DIFG for phonology and VIFG for semantics. Greater DIFG activation was associated with conflicting tonal information between words for the auditory rhyming task, suggesting this region’s role in strategic phonological processing, and greater VIFG activation was correlated with lower association between words for both the auditory and the visual meaning task, suggesting this region’s role in retrieval and selection of semantic representations. The modality- and task-specific effects in Chinese revealed by this study are similar to those found in alphabetical languages. Unlike English, we found that MFG was both modality- and task-specific, suggesting that MFG may be responsible for the visuospatial analysis of Chinese characters and orthography-to-phonology integration at a syllabic level.

Top-down control processes are thought to interact with bottom-up stimulus-driven task demands to facilitate the smooth execution of behaviour. Frontal and midline areas are believed to subserve these control processes but their distinct roles and the interactions between them remain to be fully elucidated. In this fMRI study, we utilised a GO/NO-GO task with cued and uncued inhibitory events to investigate the effect of cue-induced levels of top-down control on NO-GO trial response conflict. We found that on a within-subjects, trial-for-trial basis, high levels of top-down control, as indexed by left dorsolateral prefrontal activation prior to the NO-GO, resulted in lower levels of activation on the NO-GO trial in the pre-supplementary motor area. These results suggest that prefrontal and midline regions work together to implement cognitive control and reveal that intra-subject variability is reflected in these lateral and midline interactions.

Attentional mechanisms are a crucial prerequisite to organize behavior. Most situations may be characterized by a ‘competition’ between salient, but irrelevant stimuli and less salient, relevant stimuli. In such situations top-down and bottom-up mechanisms interact with each other. In the present fMRI study, we examined how interindividual differences in resolving situations of perceptual conflict are reflected in brain networks mediating attentional selection. Doing so, we employed a change detection task in which subjects had to detect luminance changes in the presence and absence of competing distractors. The results show that good performers presented increased activation in the orbitofrontal cortex (BA 11), anterior cingulate (BA 25), inferior parietal lobule (BA 40) and visual areas V2 and V3 but decreased activation in BA 39. This suggests that areas mediating top-down attentional control are stronger activated in this group. Increased activity in visual areas reflects distinct neuronal enhancement relating to selective attentional mechanisms in order to solve the perceptual conflict. Opposed to good performers, brain areas activated by poor performers comprised the left inferior parietal lobule (BA 39) and fronto-parietal and visual regions were continuously deactivated, suggesting that poor performers perceive stronger conflict than good performers. Moreover, the suppression of neural activation in visual areas might indicate a strategy of poor performers to inhibit the processing of the irrelevant non-target feature. These results indicate that high sensitivity in perceptual areas and increased attentional control led to less conflict in stimulus processing and consequently to higher performance in competitive attentional selection.

The segregation between cortical pathways for the identification and localization of objects is thought of as a general organizational principle in the brain. Yet, little is known about the unimodal versus multimodal nature of these processing streams. The main purpose of the present study was to test whether the auditory and tactile dual pathways converged into specialized multisensory brain areas. We used functional magnetic resonance imaging (fMRI) to compare directly in the same subjects the brain activation related to localization and identification of comparable auditory and vibrotactile stimuli. Results indicate that the right inferior frontal gyrus (IFG) and both left and right insula were more activated during identification conditions than during localization in both touch and audition. The reverse dissociation was found for the left and right inferior parietal lobules (IPL), the left superior parietal lobule (SPL) and the right precuneus-SPL, which were all more activated during localization conditions in the two modalities. We propose that specialized areas in the right IFG and the left and right insula are multisensory operators for the processing of stimulus identity whereas parts of the left and right IPL and SPL are specialized for the processing of spatial attributes independently of sensory modality.

The ability to comprehend narratives constitutes an important component of human development and experience. The neural correlates of auditory narrative comprehension in children were investigated in a large-scale functional magnetic resonance imaging (fMRI) study involving 313 subjects ages 5–18. Using group Independent Component Analysis (ICA), bilateral task-related components were found comprising the primary auditory cortex, the mid-superior temporal gyrus, the hippocampus, the angular gyrus and medial aspect of the parietal lobule (precuneus/posterior cingulate). In addition, a right-lateralized component was found involving the most posterior aspect of the superior temporal gyrus, and a left-lateralized component was found comprising the inferior frontal gyrus (including Broca’s area), the inferior parietal lobule, and the medial temporal gyrus. Using a novel data-driven analysis technique, increased task-related activity related to age was found in the components comprising the mid-superior temporal gyrus (Wernicke’s area) and the posterior aspect of the superior temporal gyrus, while decreased activity related to age was found in the component comprising the angular gyrus. The results are discussed in light of recent hypotheses involving the functional segregation of Wernicke’s area and the specific role of the mid-superior temporal gyrus in speech comprehension.

The present study examined the neural correlates of evaluations of both lying and truth-telling in different social contexts using fMRI methodology. The results demonstrated the differentiation between lying and truth-telling and between different types of lying in a network of brain regions. These regions included bilateral superior frontal gyrus (SFG), bilateral inferior parietal lobule (IPL), bilateral cuneus, right lingual gyrus (LG), right precuneus, and left postcentral gyrus (PoCG). Additionally, we found that activations in the right LG, the left IPL and the left PoCG were correlated with the off-line evaluations of truthful and untruthful communications about good and bad acts in different social contexts. These results suggest that the judgments of lying and truth-telling involving a third party might not be emotion-arousing but involve rational processing. This study is among the first to demonstrate that evaluations of truthful and untruthful communications in different social contexts can be differentiated in terms of brain BOLD (blood-oxygen-level dependent) activities.

The brain uses context and prior knowledge to repair degraded sensory inputs and improve perception. For example, listeners hear speech continuing uninterrupted through brief noises, even if the speech signal is artificially removed from the noisy epochs. In a functional MRI study, we show that this temporal filling-in process is based on two dissociable neural mechanisms: the subjective experience of illusory continuity, and the sensory repair mechanisms that support it. Areas mediating illusory continuity include the left posterior angular gyrus (AG) and superior temporal sulcus (STS) and the right STS. Unconscious sensory repair occurs in Broca’s area, bilateral anterior insula, and pre-supplementary motor area. The left AG/STS and all the repair regions show evidence for word-level template matching and communicate more when fewer acoustic cues are available. These results support a two-path process where the brain creates coherent perceptual objects by applying prior knowledge and filling-in corrupted sensory information.

In realistic auditory environments, people rely on both attentional control and attentional selection to extract intelligible signals from a cluttered background. We used functional magnetic resonance imaging to examine auditory attention to natural speech under such high processing-load conditions. Participants attended to a single talker in a group of 3, identified by the target talker's pitch or spatial location. A catch-trial design allowed us to distinguish activity due to top-down control of attention versus attentional selection of bottom-up information in both the spatial and spectral (pitch) feature domains. For attentional control, we found a left-dominant fronto-parietal network with a bias toward spatial processing in dorsal precentral sulcus and superior parietal lobule, and a bias toward pitch in inferior frontal gyrus. During selection of the talker, attention modulated activity in left intraparietal sulcus when using talker location and in bilateral but right-dominant superior temporal sulcus when using talker pitch. We argue that these networks represent the sources and targets of selective attention in rich auditory environments.

The objective of this study was to examine the neural correlates of phonological inconsistency (relationship of spelling to sound) and orthographic inconsistency (relationship of sound to spelling) in visual word processing using functional magnetic resonance imaging (fMRI). Children (9- to 15-year-old) performed a rhyming and spelling task in which two words were presented sequentially in the visual modality. Consistent with previous studies in adults, higher phonological inconsistency was associated with greater activation in several regions including left inferior frontal gyrus and medial frontal gyrus/anterior cingulate cortex. We additionally demonstrated an effect of orthographic inconsistency in these same areas, suggesting that these regions are involved in the integration of orthographic and phonological information and, with respect to the medial frontal/anterior cingulate, greater demands on executive function. Higher phonological and orthographic consistency was associated with greater activation in precuneus/posterior cingulate cortex, the putative steady state system active during resting, suggesting lower demands on cognitive resources for consistent items. Both consistency effects were larger for the rhyming compared with the spelling task suggesting greater demands of integrating spelling and sound in the former task. Finally, accuracy on the rhyming task was negatively correlated with the consistency effect in left fusiform gyrus. In particular, this region showed insensitivity to consistency in low performers, sensitivity to inconsistency (higher activity) in moderate performers, and sensitivity to inconsistency (high activation) and to consistency (deactivation). In general, these results show that the influence of spelling-sound (and sound-spelling) correspondences on processing in fusiform gyrus develops as a function of skill.

The specific role of different parietal regions to episodic retrieval is a topic of intense debate. According to the Attention to Memory (AtoM) model, dorsal parietal cortex (DPC) mediates top–down attention processes guided by retrieval goals, whereas ventral parietal cortex (VPC) mediates bottom–up attention processes captured by the retrieval output or the retrieval cue. This model also hypothesizes that the attentional functions of DPC and VPC are similar for memory and perception. To investigate this last hypothesis, we scanned participants with event-related fMRI whereas they performed memory and perception tasks, each comprising an orienting phase (top–down attention) and a detection phase (bottom–up attention). The study yielded two main findings. First, consistent with the AtoM model, orienting-related activity for memory and perception overlapped in DPC, whereas detection-related activity for memory and perception overlapped in VPC. The DPC overlap was greater in the left intraparietal sulcus, and the VPC overlap in the left TPJ. Around overlapping areas, there were differences in the spatial distribution of memory and perception activations, which were consistent with trends reported in the literature. Second, both DPC and VPC showed stronger connectivity with medial-temporal lobe during the memory task and with visual cortex during the perception task. These findings suggest that, during memory tasks, some parietal regions mediate similar attentional control processes to those involved in perception tasks (orienting in DPC vs. detection in VPC), although on different types of information (mnemonic vs. sensory).

Verbal fluency tasks have been widely used to evaluate language and executive control processes in the human brain. FMRI studies of verbal fluency, however, have used either silent word generation (which provides no behavioral measure) or cued generation of single words in order to contend with speech-related motion artifacts. In this study, we use a recently developed paradigm design to investigate the neural correlates of verbal fluency during overt, free recall, word generation so that performance and brain activity could be evaluated under conditions that more closely mirror standard behavioral test demands. We investigated verbal fluency to both letter and category cues in order to evaluate differential involvement of specific frontal and temporal lobe sites as a function of retrieval cue type, as suggested by previous neuropsychological and neuroimaging investigations. In addition, we incorporated both a task switching manipulation and an automatic speech condition in order to modulate the demand placed on executive functions. We found greater activation in the left hemisphere during category and letter fluency tasks, and greater right hemisphere activation during automatic speech. We also found that letter and category fluency tasks were associated with differential involvement of specific regions of the frontal and temporal lobes. These findings provide converging evidence that letter and category fluency performance is dependent on partially distinct neural circuitry. They also provide strong evidence that verbal fluency can be successfully evaluated in the MR environment using overt, self-paced, responses.

We show that the affective experience of touch and the sight of touch can be modulated by cognition, and investigate in an fMRI study where top-down cognitive modulations of bottom-up somatosensory and visual processing of touch and its affective value occur in the human brain. The cognitive modulation was produced by word labels, ‘Rich moisturizing cream’ or ‘Basic cream’, while cream was being applied to the forearm, or was seen being applied to a forearm. The subjective pleasantness and richness were modulated by the word labels, as were the fMRI activations to touch in parietal cortex area 7, the insula and ventral striatum. The cognitive labels influenced the activations to the sight of touch and also the correlations with pleasantness in the pregenual cingulate/orbitofrontal cortex and ventral striatum. Further evidence of how the orbitofrontal cortex is involved in affective aspects of touch was that touch to the forearm [which has C fiber Touch (CT) afferents sensitive to light touch] compared with touch to the glabrous skin of the hand (which does not) revealed activation in the mid-orbitofrontal cortex. This is of interest as previous studies have suggested that the CT system is important in affiliative caress-like touch between individuals.

Neuroimaging studies have suggested that left inferior frontal gyrus, left inferior parietal lobule and left middle temporal gyrus are critical for semantic processing in normal children. The goal of the present functional magnetic resonance imaging (fMRI) study was to determine whether these regions are systematically related to semantic processing in children (9- to 15-year-old) diagnosed with reading disorders (RD). Semantic judgments required participants to indicate whether two words were related in meaning. The strength of semantic association varied continuously from higher association pairs (e.g., king–queen) to lower association pairs (e.g. net–ship). We found that the correlation between association strength and activation was significantly weaker for RD children compared to controls in left middle temporal gyrus and left inferior parietal lobule for both the auditory and the visual modalities and in left inferior frontal gyrus for the visual modality. These results suggest that the RD children have abnormalities in semantic search/retrieval in the inferior frontal gyrus, integration of semantic information in the inferior parietal lobule and semantic lexical representations in the middle temporal gyrus. These deficits appear to be general to the semantic system and independent of modality.

Functional magnetic resonance imaging (fMRI) with continuous arterial spin labeling (CASL) was employed to monitor brain activation during narrative production of a semi-structured speech sample in healthy young adults. Subjects were asked to describe a wordless children’s picture story. Significant activations were found in bilateral prefrontal and left temporal-parietal regions during narrative production relative to description of a single picture and relative to viewing the wordless picture story while producing a nonsense word. We conclude that an inferior frontal cortex serves as a top-down organizational resource for narrative comprehension and demonstrate the feasibility of collecting extended speech samples using CASL perfusion fMRI.

The purpose of this study was to examine the neurocognitive network for processing visual word forms in native Chinese speakers using functional magnetic resonance imaging (fMRI). In order to compare the processing of phonological and semantic representations, we developed parallel rhyming and meaning association judgment tasks that required explicit access and manipulation of these representations. Subjects showed activation in left inferior/middle frontal gyri, bilateral medial frontal gyri, bilateral middle occipital/fusiform gyri, and bilateral cerebella for both the rhyming and meaning tasks. A direct comparison of the tasks revealed that the rhyming task showed more activation in the posterior dorsal region of the inferior/middle frontal gyrus (BA 9/44) and in the inferior parietal lobule (BA 40). The meaning task showed more activation in the anterior ventral region of the inferior/middle frontal gyrus (BA 47) and in the superior/middle temporal gyrus (BA 22,21). These findings are consistent with previous studies in English that suggest specialization of inferior frontal regions for the access and manipulation of phonological vs. semantic representations, but also suggest that this specialization extends to the middle frontal gyrus for Chinese. These findings are also consistent with the suggestion that the left middle temporal gyrus is involved in representing semantic information and the left inferior parietal lobule is involved in mapping between orthographic and phonological representations.

Activity within fronto-striato-temporal regions during processing of unattended auditory deviant tones and an auditory target detection task was investigated using event-related functional magnetic resonance imaging. Activation within the middle frontal gyrus, inferior frontal gyrus, anterior cingulate gyrus, superior temporal gyrus, thalamus, and basal ganglia were analyzed for differences in activity patterns between the two stimulus conditions. Unattended deviant tones elicited robust activation in the superior temporal gyrus; by contrast, attended tones evoked stronger superior temporal gyrus activation and greater frontal and striatal activation. The results suggest that attention enhances neural activation evoked by auditory pitch deviance in auditory brain regions, possibly through top-down control from the dorsolateral prefrontal cortex involved in goal-directed selection and response generation.

Functional magnetic resonance imaging (fMRI) was used to explore the neural correlates of semantic judgments to visual words in a group of 9- to 15-year-old children. Subjects were asked to indicate if word pairs were related in meaning. Consistent with previous findings in adults, children showed activation in bilateral inferior frontal gyri (Brodmann area [BA] 47, 45) and left middle temporal gyrus (BA 21). Words with strong semantic association elicited significantly greater activation in bilateral inferior parietal lobules (BA 40), suggesting stronger integration of highly related semantic features. By contrast, words with weak semantic association elicited greater activation in left inferior frontal gyrus (BA 45) and middle temporal gyrus (BA 21), suggesting more difficult feature search and more extensive access to semantic representations. We also examined whether age and skill explained unique variance in the patterns of activation. Increasing age was correlated with greater activation in left middle temporal gyrus (BA 21) and inferior parietal lobule (BA 40), suggesting that older children have more elaborated semantic representations and more complete semantic integration processes, respectively. Decreasing age was correlated with activation in right superior temporal gyrus (BA 22) and decreasing accuracy was correlated with activation in right middle temporal gyrus (BA 21), suggesting the engagement of ancillary systems in the right hemisphere for younger and lower-skill children.