Specialization of phonological and semantic processing in Chinese word reading

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.     

Changes of cognition and regional cerebral activity during acute hypoglycemia in normal subjects: A H215O positron emission tomographic study

Blurred vision and cognitive difficulties are prominent symptoms during acute insulin‐induced hypoglycemia. Our hypothesis was that changes in cerebral activity reflect these symptoms. Positron emission tomography (PET) with oxygen‐15‐labelled water was used to measure relative changes in regional cerebral blood flow (rCBF) as a marker of cerebral activity. Hypoglycemia was induced by intravenous insulin infusion in 19 healthy men performing two different cognitive tasks of varying complexity. The hypoglycemic stimulus [plasma glucose 2.2 mmol/liter (0.4)] produced a significant hormonal counterregulatory response. During the low cognitive load, rCBF decreased in response to hypoglycemia in a large bilateral area in the posterior part of the temporal lobe, and rCBF increased bilaterally in the anterior cingulate gyrus, the right frontal gyrus, the fusiform gyrus, thalamus, and the left inferior part of the frontal gyrus. During the high cognitive load, rCBF decreased bilaterally in a large region in the posterior part of the temporal gyrus and increased in the left and right anterior cingulate gyrus, left and right frontal gyrus, right parahippocampal and lingual gyrus, and left superior temporal gyrus. Visual impairment during hypoglycemia was associated with deactivation in the ventral visual stream. The anterior cingulate gyrus was activated during hypoglycemia in a load‐dependent manner. Areas on the frontal convexity were differentially activated in response to the cognitive load during hypoglycemia. Our findings suggest that hypoglycemia induces changes in sensory processing in a cognition‐independent manner, whereas activation of areas of higher order functions is influenced by cognitive load as well as hypoglycemia. © 2009 Wiley‐Liss, Inc.     

Reading fluent speech from talking faces: typical brain networks and individual differences

Listeners are able to extract important linguistic information by viewing the talker's face-a process known as 'speechreading.' Previous studies of speechreading present small closed sets of simple words and their results indicate that visual speech processing engages a wide network of brain regions in the temporal, frontal, and parietal lobes that are likely to underlie multiple stages of the receptive language system. The present study further explored this network in a large group of subjects by presenting naturally spoken sentences which tap the richer complexities of visual speech processing. Four different baselines (blank screen, static face, nonlinguistic facial gurning, and auditory speech) enabled us to determine the hierarchy of neural processing involved in speechreading and to test the claim that visual input reliably accesses sound-based representations in the auditory cortex. In contrast to passively viewing a blank screen, the static-face condition evoked activation bilaterally across the border of the fusiform gyrus and cerebellum, and in the medial superior frontal gyrus and left precentral gyrus (p < .05, whole brain corrected). With the static face as baseline, the gurning face evoked bilateral activation in the motion-sensitive region of the occipital cortex, whereas visual speech additionally engaged the middle temporal gyrus, inferior and middle frontal gyri, and the inferior parietal lobe, particularly in the left hemisphere. These latter regions are implicated in lexical stages of spoken language processing. Although auditory speech generated extensive bilateral activation across both superior and middle temporal gyri, the group-averaged pattern of speechreading activation failed to include any auditory regions along the superior temporal gyrus, suggesting that f luent visual speech does not always involve sound-based coding of the visual input. An important finding from the individual subject analyses was that activation in the superior temporal gyrus did reach significance (p < .001, small-volume corrected) for a subset of the group. Moreover, the extent of the left-sided superior temporal gyrus activity was strongly correlated with speechreading performance. Skilled speechreading was also associated with activations and deactivations in other brain regions, suggesting that individual differences ref lect the efficiency of a circuit linking sensory, perceptual, memory, cognitive, and linguistic processes rather than the operation of a single component process.     

Neural pathways involved in the processing of concrete and abstract words.

The purpose of this study was to delineate the neural pathways involved in processing concrete and abstract words using functional magnetic resonance imaging (fMRI). Word and pseudoword stimuli were presented visually, one at a time, and the participant was required to make a lexical decision. Lexical decision epochs alternated with a resting baseline. In each lexical decision epoch, the stimuli were either concrete words and pseudowords, or abstract words and pseudowords. Behavioral data indicated that, as with previous research, concrete word stimuli were processed more efficiently than abstract word stimuli. Analysis of the fMRI data indicated that processing of word stimuli, compared to the baseline condition, was associated with neural activation in the bilateral fusiform gyrus, anterior cingulate, left middle temporal gyrus, right posterior superior temporal gyrus, and left and right inferior frontal gyrus. A direct comparison between the abstract and concrete stimuli epochs yielded a significant area of activation in the right anterior temporal cortex. The results are consistent with recent positron emission tomography work showing right hemisphere activation during processing of abstract representations of language. The results are interpreted as support for a right hemisphere neural pathway in the processing of abstract word representations.     

FDG-PET mapping the brain substrates of visuo-constructive processing in Alzheimer´s disease

The anatomical basis of visuo-constructive impairment in AD is widely unexplored. FDG-PET can be used to determine functional neuronal networks underlying specific cognitive performance in the human brain. In the present study, we determined the pattern of cortical metabolism that was associated with visuo-constructive performance in AD. We employed two widely used visuo-constructive tests that differ in their demand on visual perception and processing capacity. Resting state FDG-PET scans were obtained in 29 probable AD patients, and cognitive tests were administered. We made a voxel-based regression analysis of FDG uptake to scores in visual test performance, using the SPM5 software. Performance in the CERAD Drawing test correlated with FDG uptake in the bilateral inferior temporal gyri, bilateral precuneus, right cuneus, right supramarginal gyrus and right middle temporal gyrus covering areas of dorsal and ventral visual streams. In contrast, performance in the more complex RBANS Figure Copy test correlated with FDG uptake in the bilateral fusiform gyri, right inferior temporal gyrus, left anterior cingulate gyrus, left parahippocampal gyrus, right middle temporal gyrus and right insula, encompassing the ventral visual stream and areas of higher-level visual processing. The study revealed neuronal networks underlying impaired visual test performance in AD. The extent of involvement of visual and higher order association cortex increased with greater test complexity. From a clinical point of view, both of these widely used visual tests evaluate the integrity of complementary cortical networks and may contribute complementary information on the integrity of visual processing in AD.     

PET study of the human foveal fixation system.

Positron emission tomography (PET) was used to investigate the functional anatomy of the foveal fixation system in 10 subjects scanned under three different conditions: at rest (REST), during the fixation of a central point (FIX), and while fixating the same foveal target during the presentation of peripheral visual distractors (DIS). Compared with the REST condition, both FIX and DIS tasks activated a common set of cortical areas. First, in addition to the involvement of the occipital visual cortex, both the frontal eye field (FEF) and the intraparietal sulcus (IPS) were bilaterally activated. Right frontal activation was also found in the dorsolateral prefrontal cortex, the inferior part of the precentral gyrus, and the inferior frontal gyrus. These results suggest that both FEF and IPS may constitute the main cortical regions subserving bilaterally the foveal fixation system in humans. The remaining right frontal activations may be considered as part of the anterior attentional network, supporting a role for the right frontal lobe in the allocation of the attentional mechanisms. Compared with the FIX condition, the DIS task also revealed the perceptual and cognitive processes related to the presence of peripheral visual distractors during foveal fixation. In addition to a bilateral activation of the V5/MT motion-sensitive area, a right FEF-IPS network was activated which may correspond to the engagement of the visuospatial attention. Finally, normalized regional cerebral blood flow (NrCBF) decreases were also observed during both DIS and FIX condition performance. Such NrCBF decreases were centered in the superior and middle temporal gyri, the prefrontal cortex, and the precuneus and the posterior retrosplenial part of the cingulate gyrus.     

Neuroimaging the semantic system(s)

Background: The organisation of the semantic system is controversially discussed among cognitive scientists. Whereas some authors assume that semantic processing is amodal, others propose modality-specific semantic systems. The amodal account holds that there is a single semantic system independent of the modality of stimulus presentation and nature of the concept. The multi-modal account, on the other hand, postulates that semantic knowledge is represented in modality-specific, distinct semantic systems. Attempts to integrate both accounts resulted in a hybrid model of semantic processing. Although an amodal semantic network is assumed in this view, input-dependent semantic specialisation coexists. Aims: The purpose of this study was to examine contrasting theories of the representation of the semantic knowledge with functional magnetic resonance imaging (fMRI). Methods & Procedures: Nine healthy subjects performed a semantic judgement task on graphemic words and pictures in the experimental condition, as well as a visual identity judgement task on pseudoword and three-dimensional geometric objects in the control condition. Outcomes & Results: A common activation pattern for words and pictures was found in the left inferior frontal gyrus and in the left middle temporal gyrus. Word-specific effects occurred in the left superior parietal, the left inferior occipital, the left anterior cingulate gyrus, and the right inferior frontal gyrus as well as bilaterally in the anteriomedial temporal area. Picture-specific areas involved the left supramarginal gyrus and the left cingulate gyrus. Further activation was found in the right middle frontal gyrus and bilaterally in the fusiform and inferior occipital gyrus. Conclusions: The results can best be accommodated in the framework of an integrative, hybrid model of semantic representation combining features of the amodal and the modality-specific account.     

Semantic association investigated with functional MRI and independent component analysis

Semantic association, an essential element of human language, enables discourse and inference. Neuroimaging studies have revealed localization and lateralization of semantic circuitry, making substantial contributions to cognitive neuroscience. However, because of methodological limitations, these investigations have only identified individual functional components rather than capturing the behavior of the entire network. To overcome these limitations, we have implemented group independent component analysis (ICA) to investigate the cognitive modules used by healthy adults performing the fMRI semantic decision task. When compared with the results of a standard general linear modeling (GLM) analysis, ICA detected several additional brain regions subserving semantic decision. Eight task-related group ICA maps were identified, including left inferior frontal gyrus (BA44/45), middle posterior temporal gyrus (BA39/22), angular gyrus/inferior parietal lobule (BA39/40), posterior cingulate (BA30), bilateral lingual gyrus (BA18/23), inferior frontal gyrus (L>R, BA47), hippocampus with parahippocampal gyrus (L>R, BA35/36), and anterior cingulate (BA32/24). Although most of the components were represented bilaterally, we found a single, highly left-lateralized component that included the inferior frontal gyrus and the medial and superior temporal gyri, the angular and supramarginal gyri, and the inferior parietal cortex. The presence of these spatially independent ICA components implies functional connectivity and can be equated with their modularity. These results are analyzed and presented in the framework of a biologically plausible theoretical model in preparation for similar analyses in patients with right- or left-hemispheric epilepsies.     

Brain activity related to integrative processes in visual object recognition: bottom-up integration and the modulatory influence of stored knowledge

We report evidence from a PET activation study that the inferior occipital gyri (likely to include area V2) and the posterior parts of the fusiform and inferior temporal gyri are involved in the integration of visual elements into perceptual wholes (single objects). Of these areas, the fusiform and inferior temporal gyri were more activated by tasks with recognizable stimuli than by tasks with unrecognizable stimuli. We propose that the posterior parts of the fusiform and inferior temporal gyri, compared with the inferior occipital gyri, are involved in higher level integration, due to the involvement of re-entrant activation from stored structural knowledge. Evidence in favor of this interpretation comes from the additional finding that activation of the anterior part of the left fusiform gyrus and a more anterior part of the right inferior temporal gyrus, areas previously associated with access to stored structural knowledge, was found with recognizable stimuli, but not with unrecognizable stimuli. This latter finding also indicates: (i) that subjects may not refrain from (automatically) identifying objects even if they only have to attend to the objects' global shape, and (ii) that perceptual and memorial processes can be dissociated on both functional and anatomical grounds. No evidence was obtained for the involvement of the parietal lobes in the integration of single objects.     

Lexical retrieval during fluent speech production: an fMRI study

We investigated the neural correlates of lexical retrieval during fluent speech production using fMRI. Blood oxygenation level dependent (BOLD) contrast was measured while subjects spoke about 7 Rorschach inkblots for 3 min each. Varying degrees of speech rate were elicited during each run. In a within-subject design, the number of words produced was correlated with the BOLD contrast in the two runs in each subject that showed the highest variance of speech output. The rate of articulation was positively correlated with activation in the left superior temporal (BA 22) and supramarginal (BA 39/ 40) gyri. The main negative correlations were in the fusiform gyri bilaterally (BA 19), the posterior cingulate (BA 30) and superior occipital gyrus (BA 19). Lexical retrieval during continuous language production engages areas in the left temporal and inferior parietal cortex. This pattern of activation differs from that evident during the generation of single words (verbal fluency), which is more associated with left prefrontal activation.     

The signer and the sign: Cortical correlates of person identity and language processing from point-light displays

In this study, the first to explore the cortical correlates of signed language (SL) processing under point-light display conditions, the observer identified either a signer or a lexical sign from a display in which different signers were seen producing a number of different individual signs. Many of the regions activated by point-light under these conditions replicated those previously reported for full-image displays, including regions within the inferior temporal cortex that are specialised for face and body-part identification, although such body parts were invisible in the display. Right frontal regions were also recruited – a pattern not usually seen in full-image SL processing. This activation may reflect the recruitment of information about person identity from the reduced display. A direct comparison of identify-signer and identify-sign conditions showed these tasks relied to a different extent on the posterior inferior regions. Signer identification elicited greater activation than sign identification in (bilateral) inferior temporal gyri (BA 37/19), fusiform gyri (BA 37), middle and posterior portions of the middle temporal gyri (BAs 37 and 19), and superior temporal gyri (BA 22 and 42). Right inferior frontal cortex was a further focus of differential activation (signer > sign).These findings suggest that the neural systems supporting point-light displays for the processing of SL rely on a cortical network including areas of the inferior temporal cortex specialized for face and body identification. While this might be predicted from other studies of whole body point-light actions (Vaina, Solomon, Chowdhury, Sinha, & Belliveau, 2001) it is not predicted from the perspective of spoken language processing, where voice characteristics and speech content recruit distinct cortical regions (Stevens, 2004) in addition to a common network. In this respect, our findings contrast with studies of voice/speech recognition (Von Kriegstein, Kleinschmidt, Sterzer, & Giraud, 2005). Inferior temporal regions associated with the visual recognition of a person appear to be required during SL processing, for both carrier and content information.     

Differential effects of orthographic and phonological consistency in cortex for children with and without reading impairment

One of the central challenges in mastering English is becoming sensitive to consistency from spelling to sound (i.e. phonological consistency) and from sound to spelling (i.e. orthographic consistency). Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of consistency in 9–15-year-old Normal and Impaired Readers during a rhyming task in the visual modality. In line with our previous study [Bolger, D. J., Hornickel, J., Cone, N. E., Burman, D. D., & Booth, J. R. (in press). Neural correlates of orthographic and phonological consistency effects in children. Human Brain Mapping], for Normal Readers, lower phonological and orthographic consistency were associated with greater activation in several regions including bilateral inferior/middle frontal gyri, bilateral anterior cingulate cortex as well as left fusiform gyrus. Impaired Readers activated only bilateral anterior cingulate cortex in response to decreasing consistency. Group comparisons revealed that, relative to Impaired Readers, Normal Readers exhibited a larger response in this network for lower phonological consistency whereas orthographic consistency differences were limited. Lastly, brain–behavior correlations revealed a significant relationship between skill (i.e. Phonological Awareness and non-word decoding) and cortical consistency effects for Impaired Readers in left inferior/middle frontal gyri and left fusiform gyrus. Impaired Readers with higher skill showed greater activation for higher consistency. This relationship was reliably different from that of Normal Readers in which higher skill was associated with greater activation for lower consistency. According to single-route or connectionist models, these results suggest that Impaired Readers with higher skill devote neural resources to representing the mapping between orthography and phonology for higher consistency words, and therefore do not robustly activate this network for lower consistency words.     

Neural correlates of episodic and semantic memory retrieval in borderline personality disorder: An fMRI study

Verbal memory impairment in borderline personality disorder (BPD) is still a matter of debate. In this study we combine investigations of both, memory retrieval as well as underlying neural circuits in BPD. Functional magnetic resonance imaging (fMRI) was used to study regional brain activation in 18 right-handed female patients with BPD and 18 matched controls during the retrieval of an episodic memory retrieval (EMR) task (free recall of a word list) and a semantic memory retrieval (SMR) task (verbal fluency). Despite unaffected performance in EMR and SMR, patients with BPD showed task-specific increased activation compared with controls. During EMR, the increased activation encompassed the posterior cingulate cortex bilaterally, the left middle and superior temporal gyrus, the right inferior frontal gyrus, and the right angular gyrus. SMR was associated with increased activation of the posterior cingulate cortex, of the right fusiform gyrus, of the left anterior cingulate cortex, and of the left postcentral gyrus. Our findings suggest that BPD patients may need to engage larger brain areas to reach a level of performance in episodic and semantic retrieval tasks that is comparable to that of healthy controls.     

Modality independence of word comprehension

Functional magnetic resonance imaging (fMRI) was used to examine the functional anatomy of word comprehension in the auditory and visual modalities of presentation. We asked our subjects to determine if word pairs were semantically associated (e.g., table, chair) and compared this to a reference task where they were asked to judge whether word pairs rhymed (e.g., bank, tank). This comparison showed task-specific and modality-independent activation for semantic processing in the heteromodal cortices of the left inferior frontal gyrus (BA 46, 47) and left middle temporal gyrus (BA 21). There were also modality-specific activations in the fusiform gyrus (BA 37) for written words and in the superior temporal gyrus (BA 22) for spoken words. Our findings are consistent with the hypothesis that word form recognition (lexical encoding) occurs in unimodal cortices and that heteromodal brain regions in the anterior as well as posterior components of the language network subserve word comprehension (semantic decoding).     

An fMRI study of sex differences in brain activation during object naming

We present a neuroimaging experiment that examines whether males and females use distinct brain systems while performing a confrontational naming task, with specific attention to the possibility of laterality differences, as suggested by some theories of sex differences in language processing. We further address whether sex-based differences in functional brain organization might interact with object category distinctions, given that previous behavioral studies have shown some consistent processing differences between the sexes with respect to tools versus plants. Functional magnetic resonance imaging (fMRI) data were collected from 26 participants (13 males and 13 females). Main effect and interaction analyses reveal no discernable laterality differences between the sexes. All other results, however, were consistent with previous object-naming studies. Global effects revealed dominant foci in fusiform gyrus, left posterior middle temporal gyrus, left basal ganglia/thalamus, left middle/inferior frontal gyri, left frontal operculum, left supplementary motor area/dorsal anterior cingulate, and left pre-central gyrus. Main contrasts for tools versus plants were likewise consistent with previous fMRI studies. Although men and women showed no discernable activation differences, hemispheric or otherwise, when collapsed across object categories, sex-by-category analyses showed selective activation for females in dorsal anterior cingulate gyrus and left posterior middle temporal gyrus for tools, and selective activation for males in left posterior middle temporal gyrus for plants. We discuss the relevance of these sex-by-category effects to previous behavioral findings and theories that relate to vocabulary differences between the sexes.     

Temporal lobe abnormalities in semantic processing by criminal psychopaths as revealed by functional magnetic resonance imaging

We tested the hypothesis that psychopathy is associated with abnormalities in semantic processing of linguistic information. Functional magnetic resonance imaging (fMRI) was used to elucidate and characterize the neural architecture underlying lexico-semantic processes in criminal psychopathic individuals and in a group of matched control participants. Participants performed a lexical decision task in which blocks of linguistic stimuli alternated with a resting baseline condition. In each lexical decision block, the stimuli were either concrete words and pseudowords or abstract words and pseudowords. Consistent with our hypothesis, psychopathic individuals, relative to controls, showed poorer behavioral performance for processing abstract words. Analysis of the fMRI data for both groups indicated that processing of word stimuli, compared with the resting baseline condition, was associated with neural activation in bilateral fusiform gyrus, anterior cingulate, left middle temporal gyrus, right posterior superior temporal gyrus, and left and right inferior frontal gyrus. Analyses confirmed our prediction that psychopathic individuals would fail to show the appropriate neural differentiation between abstract and concrete stimuli in the right anterior temporal gyrus and surrounding cortex. The results are consistent with other studies of semantic processing in psychopathy and support the theory that psychopathy is associated with right hemisphere abnormalities for processing conceptually abstract material.     

Temporal lobe abnormalities in semantic processing by criminal psychopaths as revealed by functional magnetic resonance imaging

We tested the hypothesis that psychopathy is associated with abnormalities in semantic processing of linguistic information. Functional magnetic resonance imaging (fMRI) was used to elucidate and characterize the neural architecture underlying lexico-semantic processes in criminal psychopathic individuals and in a group of matched control participants. Participants performed a lexical decision task in which blocks of linguistic stimuli alternated with a resting baseline condition. In each lexical decision block, the stimuli were either concrete words and pseudowords or abstract words and pseudowords. Consistent with our hypothesis, psychopathic individuals, relative to controls, showed poorer behavioral performance for processing abstract words. Analysis of the fMRI data for both groups indicated that processing of word stimuli, compared with the resting baseline condition, was associated with neural activation in bilateral fusiform gyrus, anterior cingulate, left middle temporal gyrus, right posterior superior temporal gyrus, and left and right inferior frontal gyrus. Analyses confirmed our prediction that psychopathic individuals would fail to show the appropriate neural differentiation between abstract and concrete stimuli in the right anterior temporal gyrus and surrounding cortex. The results are consistent with other studies of semantic processing in psychopathy and support the theory that psychopathy is associated with right hemisphere abnormalities for processing conceptually abstract material.     

Modality- and task-specific brain regions involved in Chinese lexical processing

fMRI was used to examine lexical processing in native adult Chinese speakers. A 2 task (semantics and phonology) x 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.