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1.  A dynamic causal modeling analysis of the effective connectivities underlying top-down letter processing 
Neuropsychologia  2011;49(5):10.1016/j.neuropsychologia.2011.01.011.
The present study employed Dynamic Causal Modeling to investigate the effective functional connectivity between regions of the neural network involved in top-down letter processing. We used an illusory letter detection paradigm in which participants detected letters while viewing pure noise images. When participants detected letters, the response of the right middle occipital gyrus (MOG) in the visual cortex was enhanced by increased feed-backward connectivity from the left inferior frontal gyrus (IFG). In addition, illusory letter detection increased feed-forward connectivity from the right MOG to the left inferior parietal lobules. Originating in the left IFG, this top-down letter processing network may facilitate the detection of letters by activating letter processing areas within the visual cortex. This activation in turns may highlight the visual features of letters and send letter information to activate the associated phonological representations in the identified parietal region.
PMCID: PMC3817006  PMID: 21237182
letter processing; word processing; top-down processing; fMRI; dynamic causal modeling
2.  Word and Letter String Processing Networks in Schizophrenia: Evidence for Anomalies and Compensation 
Brain and language  2008;107(2):158-166.
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.
PMCID: PMC2599869  PMID: 18829095
Schizophrenia and language; Lateralization; Lexical-semantic processing; Imaging; Effective Connectivity; Modeling
3.  A Common Left Occipito-Temporal Dysfunction in Developmental Dyslexia and Acquired Letter-By-Letter Reading? 
PLoS ONE  2010;5(8):e12073.
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.
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.
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.
PMCID: PMC2920311  PMID: 20711448
4.  A distributed neural system for top-down face processing 
Neuroscience letters  2008;451(1):6-10.
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.
PMCID: PMC2634849  PMID: 19121364
top-down processing; psychophysiological interaction (PPI); distributed cortical network; fMRI; face processing
5.  Neuroanatomical dissociation between bottom-up and top-down processes of visuospatial selective attention 
NeuroImage  2006;32(2):842-853.
Allocation of attentional resources to portions of the available sensory input can be regulated by bottom-up processes, i.e. spontaneous orientation towards an oncoming stimulus (stimulus-driven attention), and by top-down processes, i.e. intentionally and driven by knowledge, expectation and goals. The present study aimed at advancing the understanding of brain networks mediating bottom-up and top-down control of visuospatial attention by employing a paradigm that parametrically varied demands on these two processes. Spatial predictability of peripheral targets was parametrically varied by centrally cueing one, two, three or four of four possible locations. Reaction time decreased linearly with more precise valid cueing of the target location and increased with more precise invalid cueing. Event-related functional magnetic resonance imaging (fMRI) enabled measurement of blood oxygenation level-dependent (BOLD) responses to cues and to targets. A mostly left-hemispheric network consisting of left intraparietal sulcus, inferior and superior parietal lobule, bilateral precuneus, middle frontal gyri including superior frontal sulci, and middle occipital gyri displayed BOLD responses to cues that increased linearly with more precise spatial cueing, indicating engagement by top-down spatial selective attention. In contrast, bilateral temporoparietal junction, cingulate gyrus, right precentral gyrus and anterior and posterior insula, bilateral fusiform gyri, lingual gyri and cuneus displayed BOLD responses to targets that increased with their spatial unpredictability, indicating engagement by stimulus-driven orienting. The results suggest two largely dissociated neural networks mediating top-down and bottom-up control of visuospatial selective attention.
PMCID: PMC2652125  PMID: 16757180
6.  Developmental Increase in Top–Down and Bottom–Up Processing in a Phonological Task: An Effective Connectivity, fMRI Study 
We examined age-related changes in the interactions among brain regions in children performing rhyming judgments on visually presented words. The difficulty of the task was manipulated by including a conflict between task-relevant (phonological) information and task-irrelevant (orthographic) information. The conflicting conditions included pairs of words that rhyme despite having different spelling patterns ( jazz–has), or words that do not rhyme despite having similar spelling patterns ( pint–mint). These were contrasted with nonconflicting pairs that have similar orthography and phonology (dime–lime) or different orthography and phonology ( press–list). Using fMRI, we examined effective connectivity among five left hemisphere regions of interest: fusiform gyrus (FG), inferior frontal gyrus (IFG), intraparietal sulcus (IPS), lateral temporal cortex (LTC), and medial frontal gyrus (MeFG). Age-related increases were observed in the influence of the IFG and FG on the LTC, but only in conflicting conditions. These results reflect a developmental increase in the convergence of bottom–up and top–down information on the LTC. In older children, top–down control process may selectively enhance the sensitivity of the LTC to bottom–up information from the FG. This may be evident especially in situations that require selective enhancement of task-relevant versus task-irrelevant information. Altogether these results provide a direct evidence for a developmental increase in top–down control processes in language processing. The developmental increase in bottom–up processing may be secondary to the enhancement of top–down processes.
PMCID: PMC2821219  PMID: 18702576
7.  Fixation-Related fMRI Analysis in the Domain of Reading Research: Using Self-Paced Eye Movements as Markers for Hemodynamic Brain Responses During Visual Letter String Processing 
Cerebral Cortex (New York, NY)  2013;24(10):2647-2656.
The present study investigated the feasibility of using self-paced eye movements during reading (measured by an eye tracker) as markers for calculating hemodynamic brain responses measured by functional magnetic resonance imaging (fMRI). Specifically, we were interested in whether the fixation-related fMRI analysis approach was sensitive enough to detect activation differences between reading material (words and pseudowords) and nonreading material (line and unfamiliar Hebrew strings). Reliable reading-related activation was identified in left hemisphere superior temporal, middle temporal, and occipito-temporal regions including the visual word form area (VWFA). The results of the present study are encouraging insofar as fixation-related analysis could be used in future fMRI studies to clarify some of the inconsistent findings in the literature regarding the VWFA. Our study is the first step in investigating specific visual word recognition processes during self-paced natural sentence reading via simultaneous eye tracking and fMRI, thus aiming at an ecologically valid measurement of reading processes. We provided the proof of concept and methodological framework for the analysis of fixation-related fMRI activation in the domain of reading research.
PMCID: PMC4153805  PMID: 23645718
cerebrum; functional magnetic resonance imaging; language; visual word form area; visual word recognition
8.  On the functional neuroanatomy of visual word processing: Effects of case and letter deviance 
Journal of cognitive neuroscience  2009;21(2):222-229.
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.
PMCID: PMC2976854  PMID: 18476755
Functional MRI; visual word recognition; occipitotemporal cortex; visual word form area; orthographic processing
9.  Faces in the mist: illusory face and letter detection 
i-Perception  2011;2(5):458-476.
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.
PMCID: PMC3485785  PMID: 23145238
vision; face perception; reverse correlation; letter perception; top down; false detection
10.  Effective connectivities of cortical regions for top-down face processing: A Dynamic Causal Modeling study 
Brain research  2010;1340:40-51.
To study top-down face processing, the present study used an experimental paradigm in which participants detected non-existent faces in pure noise images. Conventional BOLD signal analysis identified three regions involved in this illusory face detection. These regions included the left orbitofrontal cortex (OFC) in addition to the right fusiform face area (FFA) and right occipital face area (OFA), both of which were previously known to be involved in both top-down and bottom-up processing of faces. We used Dynamic Causal Modeling (DCM) and Bayesian model selection to further analyze the data, revealing both intrinsic and modulatory effective connectivities among these three cortical regions. Specifically, our results support the claim that the orbitofrontal cortex plays a crucial role in the top-down processing of faces by regulating the activities of the occipital face area, and the occipital face area in turn detects the illusory face features in the visual stimuli and then provides this information to the fusiform face area for further analysis.
PMCID: PMC3724518  PMID: 20423709
Face processing; Top-down processing; Bottom-up processing; Dynamic Causal Modeling (DCM); Orbitofrontal cortex (OFC)
NeuroImage  2008;40(4):1807-1814.
Although it is accepted that visual cortical areas are recruited during touch, it remains uncertain whether this depends on top-down inputs mediating visual imagery or engagement of modality-independent representations by bottom-up somatosensory inputs. Here we addressed this by examining effective connectivity in humans during haptic perception of shape and texture with the right hand. Multivariate Granger causality analysis of functional magnetic resonance imaging (fMRI) data was conducted on a network of regions that were shape- or texture-selective. A novel network reduction procedure was employed to eliminate connections that did not contribute significantly to overall connectivity. Effective connectivity during haptic perception was found to involve a variety of interactions between areas generally regarded as somatosensory, multisensory, visual and motor, emphasizing flexible cooperation between different brain regions rather than rigid functional separation. The left postcentral sulcus (PCS), left precentral gyrus and right posterior insula were important sources of connections in the network. Bottom-up somatosensory inputs from the left PCS and right posterior insula fed into visual cortical areas, both the shape-selective right lateral occipital complex (LOC) and the texture-selective right medial occipital cortex (probable V2). In addition, top-down inputs from left postero-supero-medial parietal cortex influenced the right LOC. Thus, there is strong evidence for the bottom-up somatosensory inputs predicted by models of visual cortical areas as multisensory processors and suggestive evidence for top-down parietal (but not prefrontal) inputs that could mediate visual imagery. This is consistent with modality-independent representations accessible through both bottom-up sensory inputs and top-down processes such as visual imagery.
PMCID: PMC2483676  PMID: 18329290
12.  Letter representations in writing: an fMRI adaptation approach 
Behavioral and neuropsychological research in reading and spelling has provided evidence for the role of the following types of orthographic representations in letter writing: letter shapes, letter case, and abstract letter identities. We report on the results of an fMRI investigation designed to identify the neural substrates of these different representational types. Using an fMRI adaptation paradigm we examined the neural distribution of inhibition and release from inhibition in a letter-writing task in which, on every trial, participants produced three repetitions of the same letter and a fourth letter that was either identical to (no-change trial) or different from the previous three (change trial). Change trials involved a change in the shape, case, and/or identity of the letter. After delineating the general letter writing network by identifying areas that exhibited significant neural adaptation effects on no-change trials, we used deconvolution analysis to examine this network for effects of release from inhibition on change trials. In this way we identified regions specifically associated with the representation of letter shape (in the left SFS and SFG/pre-CG) and letter identity [in the left fusiform gyrus (FG)] or both [right cerebellum, left post-central gyrus (post-CG), and left middle frontal gyrus (MFG)]. No regions were associated with the representation of letter case. This study showcases an investigational approach that allows for the differentiation of the neurotopography of the representational types that are key to our ability to produce written language.
PMCID: PMC3809555  PMID: 24194724
letter; writing; fMRI; fMRI adaptation; neural habituation; letter shape; letter identity; letter case
13.  Emotional face expression modulates occipital-frontal effective connectivity during memory formation in a bottom-up fashion 
This study investigated the role of bottom-up and top-down neural mechanisms in the processing of emotional face expression during memory formation. Functional brain imaging data was acquired during incidental learning of positive (“happy”), neutral and negative (“angry” or “fearful”) faces. Dynamic Causal Modeling (DCM) was applied on the functional magnetic resonance imaging (fMRI) data to characterize effective connectivity within a brain network involving face perception (inferior occipital gyrus and fusiform gyrus) and successful memory formation related areas (hippocampus, superior parietal lobule, amygdala, and orbitofrontal cortex). The bottom-up models assumed processing of emotional face expression along feed forward pathways to the orbitofrontal cortex. The top-down models assumed that the orbitofrontal cortex processed emotional valence and mediated connections to the hippocampus. A subsequent recognition memory test showed an effect of negative emotion on the response bias, but not on memory performance. Our DCM findings showed that the bottom-up model family of effective connectivity best explained the data across all subjects and specified that emotion affected most bottom-up connections to the orbitofrontal cortex, especially from the occipital visual cortex and superior parietal lobule. Of those pathways to the orbitofrontal cortex the connection from the inferior occipital gyrus correlated with memory performance independently of valence. We suggest that bottom-up neural mechanisms support effects of emotional face expression and memory formation in a parallel and partially overlapping fashion.
PMCID: PMC4407577  PMID: 25954169
Dynamic Causal Modeling; fMRI; facial affect; memory formation
14.  Neural Dissociation of Number from Letter Recognition and Its Relationship to Parietal Numerical Processing 
The visual recognition of letters dissociates from the recognition of numbers at both the behavioral and neural level. In this article, using fMRI, we investigate whether the visual recognition of numbers dissociates from letters, thereby establishing a double dissociation. In Experiment 1, participants viewed strings of consonants and Arabic numerals. We found that letters activated the left midfusiform and inferior temporal gyri more than numbers, replicating previous studies, whereas numbers activated a right lateral occipital area more than letters at the group level. Because the distinction between letters and numbers is culturally defined and relatively arbitrary, this double dissociation provides some of the strongest evidence to date that a neural dissociation can emerge as a result of experience. We then investigated a potential source of the observed neural dissociation. Specifically, we tested the hypothesis that lateralization of visual number recognition depends on lateralization of higher-order numerical processing. In Experiment 2, the same participants performed addition, subtraction, and counting on arrays of nonsymbolic stimuli varying in numerosity, which produced neural activity in and around the intraparietal sulcus, a region associated with higher-order numerical processing. We found that individual differences in the lateralization of number activity in visual cortex could be explained by individual differences in the lateralization of numerical processing in parietal cortex, suggesting a functional relationship between the two regions. Together, these results demonstrate a neural double dissociation between letter and number recognition and suggest that higher-level numerical processing in parietal cortex may influence the neural organization of number processing in visual cortex.
PMCID: PMC3357212  PMID: 21736455
15.  The Temporal Dynamics of Implicit Processing of Non-Letter, Letter, and Word-Forms in the Human Visual Cortex 
The decoding of visually presented line segments into letters, and letters into words, is critical to fluent reading abilities. Here we investigate the temporal dynamics of visual orthographic processes, focusing specifically on right hemisphere contributions and interactions between the hemispheres involved in the implicit processing of visually presented words, consonants, false fonts, and symbolic strings. High-density EEG was recorded while participants detected infrequent, simple, perceptual targets (dot strings) embedded amongst a of character strings. Beginning at 130 ms, orthographic and non-orthographic stimuli were distinguished by a sequence of ERP effects over occipital recording sites. These early latency occipital effects were dominated by enhanced right-sided negative-polarity activation for non-orthographic stimuli that peaked at around 180 ms. This right-sided effect was followed by bilateral positive occipital activity for false-fonts, but not symbol strings. Moreover the size of components of this later positive occipital wave was inversely correlated with the right-sided ROcc180 wave, suggesting that subjects who had larger early right-sided activation for non-orthographic stimuli had less need for more extended bilateral (e.g., interhemispheric) processing of those stimuli shortly later. Additional early (130–150 ms) negative-polarity activity over left occipital cortex and longer-latency centrally distributed responses (>300 ms) were present, likely reflecting implicit activation of the previously reported ‘visual-word-form’ area and N400-related responses, respectively. Collectively, these results provide a close look at some relatively unexplored portions of the temporal flow of information processing in the brain related to the implicit processing of potentially linguistic information and provide valuable information about the interactions between hemispheres supporting visual orthographic processing.
PMCID: PMC2796900  PMID: 20046826
word reading; ERPs; visual cortex; visual orthography
16.  Developmental changes in category-specific brain responses to numbers and letters in a working memory task 
NeuroImage  2008;44(4):1404-1414.
Neuroimaging studies have identified a common network of brain regions involving the prefrontal and parietal cortices across a variety of working memory (WM) tasks. However, previous studies have also reported category-specific dissociations of activation within this network. In this study, we investigated the development of category-specific activation in a WM task with digits, letters, and faces. Eight-year-old children and adults performed a 2-back WM task while their brain activity was measured using functional magnetic resonance imaging (fMRI). Overall, children were significantly slower and less accurate than adults on all three WM conditions (digits, letters, and faces); however, within each age group, behavioral performance across the three conditions was very similar. FMRI results revealed category-specific activation in adults but not children in the intraparietal sulcus for the digit condition. Likewise, during the letter condition, category-specific activation was observed in adults but not children in the left occipital–temporal cortex. In contrast, children and adults showed highly similar brain-activity patterns in the lateral fusiform gyri when solving the 2-back WM task with face stimuli. Our results suggest that 8-year-old children do not yet engage the typical brain regions that have been associated with abstract or semantic processing of numerical symbols and letters when these processes are task-irrelevant and the primary task is demanding. Nevertheless, brain activity in letter-responsive areas predicted children’s spelling performance underscoring the relationship between abstract processing of letters and linguistic abilities. Lastly, behavioral performance on the WM task was predictive of math and language abilities highlighting the connection between WM and other cognitive abilities in development.
PMCID: PMC2659412  PMID: 19027079
17.  Repetition of letter strings leads to activation of and connectivity with word-related regions 
Neuroimage  2011;59(3):2839-2849.
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.
PMCID: PMC3254793  PMID: 21982931
letter strings; fMRI; connectivity; reading; learning; plasticity
18.  Children with Reading Disability Show Brain Differences in Effective Connectivity for Visual, but Not Auditory Word Comprehension 
PLoS ONE  2010;5(10):e13492.
Previous literature suggests that those with reading disability (RD) have more pronounced deficits during semantic processing in reading as compared to listening comprehension. This discrepancy has been supported by recent neuroimaging studies showing abnormal activity in RD during semantic processing in the visual but not in the auditory modality. Whether effective connectivity between brain regions in RD could also show this pattern of discrepancy has not been investigated.
Methodology/Principal Findings
Children (8- to 14-year-olds) were given a semantic task in the visual and auditory modality that required an association judgment as to whether two sequentially presented words were associated. Effective connectivity was investigated using Dynamic Causal Modeling (DCM) on functional magnetic resonance imaging (fMRI) data. Bayesian Model Selection (BMS) was used separately for each modality to find a winning family of DCM models separately for typically developing (TD) and RD children. BMS yielded the same winning family with modulatory effects on bottom-up connections from the input regions to middle temporal gyrus (MTG) and inferior frontal gyrus(IFG) with inconclusive evidence regarding top-down modulations. Bayesian Model Averaging (BMA) was thus conducted across models in this winning family and compared across groups. The bottom-up effect from the fusiform gyrus (FG) to MTG rather than the top-down effect from IFG to MTG was stronger in TD compared to RD for the visual modality. The stronger bottom-up influence in TD was only evident for related word pairs but not for unrelated pairs. No group differences were noted in the auditory modality.
This study revealed a modality-specific deficit for children with RD in bottom-up effective connectivity from orthographic to semantic processing regions. There were no group differences in connectivity from frontal regions, suggesting that the core deficit in RD is not in top-down modulation.
PMCID: PMC2963599  PMID: 21049093
19.  Neural Correlates of Individual Performance Differences in Resolving Perceptual Conflict 
PLoS ONE  2012;7(8):e42849.
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.
PMCID: PMC3423441  PMID: 22916169
20.  Neural systems supporting lexical search guided by letter and semantic category cues: A self-paced overt response fMRI study of verbal fluency 
NeuroImage  2009;49(1):1099-1107.
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.
PMCID: PMC2832834  PMID: 19632335
21.  Sensory-to-motor integration during auditory repetition: a combined fMRI and lesion study 
The aim of this paper was to investigate the neurological underpinnings of auditory-to-motor translation during auditory repetition of unfamiliar pseudowords. We tested two different hypotheses. First we used functional magnetic resonance imaging in 25 healthy subjects to determine whether a functionally defined area in the left temporo-parietal junction (TPJ), referred to as Sylvian-parietal-temporal region (Spt), reflected the demands on auditory-to-motor integration during the repetition of pseudowords relative to a semantically mediated nonverbal sound-naming task. The experiment also allowed us to test alternative accounts of Spt function, namely that Spt is involved in subvocal articulation or auditory processing that can be driven either bottom-up or top-down. The results did not provide convincing evidence that activation increased in either Spt or any other cortical area when non-semantic auditory inputs were being translated into motor outputs. Instead, the results were most consistent with Spt responding to bottom up or top down auditory processing, independent of the demands on auditory-to-motor integration. Second, we investigated the lesion sites in eight patients who had selective difficulties repeating heard words but with preserved word comprehension, picture naming and verbal fluency (i.e., conduction aphasia). All eight patients had white-matter tract damage in the vicinity of the arcuate fasciculus and only one of the eight patients had additional damage to the Spt region, defined functionally in our fMRI data. Our results are therefore most consistent with the neurological tradition that emphasizes the importance of the arcuate fasciculus in the non-semantic integration of auditory and motor speech processing.
PMCID: PMC3908611  PMID: 24550807
fMRI; lesions; language; speech; aphasia
22.  Neural Correlates of Letter Reversal in Children and Adults 
PLoS ONE  2014;9(5):e98386.
Children often make letter reversal errors when first learning to read and write, even for letters whose reversed forms do not appear in normal print. However, the brain basis of such letter reversal in children learning to read is unknown. The present study compared the neuroanatomical correlates (via functional magnetic resonance imaging) and the electrophysiological correlates (via event-related potentials or ERPs) of this phenomenon in children, ages 5–12, relative to young adults. When viewing reversed letters relative to typically oriented letters, adults exhibited widespread occipital, parietal, and temporal lobe activations, including activation in the functionally localized visual word form area (VWFA) in left occipito-temporal cortex. Adults exhibited significantly greater activation than children in all of these regions; children only exhibited such activation in a limited frontal region. Similarly, on the P1 and N170 ERP components, adults exhibited significantly greater differences between typical and reversed letters than children, who failed to exhibit significant differences between typical and reversed letters. These findings indicate that adults distinguish typical and reversed letters in the early stages of specialized brain processing of print, but that children do not recognize this distinction during the early stages of processing. Specialized brain processes responsible for early stages of letter perception that distinguish between typical and reversed letters may develop slowly and remain immature even in older children who no longer produce letter reversals in their writing.
PMCID: PMC4032318  PMID: 24859328
23.  Attentional control during the transient updating of cue information 
Brain research  2008;1247:149-158.
The goal of the present study was to investigate the neural correlates of top-down control of switching behavior in humans and to contrast them to those observed during switching behavior guided by bottom-up mechanisms. In the main experimental condition (color-cue), which was guided by top-down control, a central cue indicated the color of a peripheral grating on which the subject performed an orientation judgment. For switch trials, the color of the cue on the current trial was different from the color on the previous trial. For non-switch trials, the color of the cue on the current trial was the same as the color in the preceding trial. During a control condition (pop-out), which was guided by bottom-up saliency, the target grating was defined by color contrast; again both switch and non-switch trials occurred. We observed stronger evoked responses during the color-cue task relative to the pop-out task in the inferior parietal lobule (IPL), frontal eye field (FEF), middle frontal gyrus (MFG), and inferior frontal gyrus (IFG). The contrast of switch vs. non-switch trials revealed activations in regions that were engaged when there was a change in the identity of the target. Collectively, switch trials evoked stronger responses relative to non-switch trials in fronto-parietal regions that appeared to be left lateralized, including left intraparietal sulcus (IPS) and left MFG/IFG. Task by trial type interactions (switch > non-switch during color-cue relative to pop-out) were observed in several fronto-parietal regions, including IPS, FEF, MFG and IFG, in addition to regions in visual cortex. Our findings suggest that, within the fronto-parietal attentional network, the IPS and MFG/IFG appear to be most heavily involved in attentive cue updating. Furthermore, several visual regions engaged by oriented gratings were strongly affected by cue updating, raising the possibility that they were the recipient of top-down signals that were generated when cue information was updated.
PMCID: PMC2661537  PMID: 18992228
24.  Rhythmic and melodic deviations in musical sequences recruit different cortical areas for mismatch detection 
The mismatch negativity (MMN), an event-related potential (ERP) representing the violation of an acoustic regularity, is considered as a pre-attentive change detection mechanism at the sensory level on the one hand and as a prediction error signal on the other hand, suggesting that bottom-up as well as top-down processes are involved in its generation. Rhythmic and melodic deviations within a musical sequence elicit a MMN in musically trained subjects, indicating that acquired musical expertise leads to better discrimination accuracy of musical material and better predictions about upcoming musical events. Expectation violations to musical material could therefore recruit neural generators that reflect top-down processes that are based on musical knowledge. We describe the neural generators of the musical MMN for rhythmic and melodic material after a short-term sensorimotor-auditory (SA) training. We compare the localization of musical MMN data from two previous MEG studies by applying beamformer analysis. One study focused on the melodic harmonic progression whereas the other study focused on rhythmic progression. The MMN to melodic deviations revealed significant right hemispheric neural activation in the superior temporal gyrus (STG), inferior frontal cortex (IFC), and the superior frontal (SFG) and orbitofrontal (OFG) gyri. IFC and SFG activation was also observed in the left hemisphere. In contrast, beamformer analysis of the data from the rhythm study revealed bilateral activation within the vicinity of auditory cortices and in the inferior parietal lobule (IPL), an area that has recently been implied in temporal processing. We conclude that different cortical networks are activated in the analysis of the temporal and the melodic content of musical material, and discuss these networks in the context of the dual-pathway model of auditory processing.
PMCID: PMC3675320  PMID: 23759929
music perception; mismatch negativity; beamforming; cortex
25.  Deficient orthographic and phonological representations in children with dyslexia revealed by brain activation patterns 
The current study examined the neuro-cognitive network of visual word rhyming judgment in 14 children with dyslexia and 14 age-matched control children (8- to 14-year-olds) using functional magnetic resonance imaging (fMRI).
In order to manipulate the difficulty of mapping orthography to phonology, we used conflicting and non-conflicting trials. The words in conflicting trials either had similar orthography but different phonology (e.g., pint-mint) or similar phonology but different orthography (e.g., jazz-has). The words in non-conflicting trials had similar orthography and phonology (e.g., gate-hate) or different orthography and phonology (e.g., press-list).
There were no differences in brain activation between the controls and children with dyslexia in the easier non-conflicting trials. However, the children with dyslexia showed less activation than the controls in left inferior frontal gyrus (BA 45/44/47/9), left inferior parietal lobule (BA 40), left inferior temporal gyrus/fusiform gyrus (BA 20/37) and left middle temporal gyrus (BA 21) for the more difficult conflicting trials. For the direct comparison of conflicting minus non-conflicting trials, controls showed greater activation than children with dyslexia in left inferior frontal gyrus (BA 9/45/46) and medial frontal gyrus (BA 8). Children with dyslexia did not show greater activation than controls for any comparison.
Reduced activation in these regions suggests that children with dyslexia have deficient orthographic representations in ventral temporal cortex as well as deficits in mapping between orthographic and phonological representations in inferior parietal cortex. The greater activation for the controls in inferior frontal gyrus could reflect more effective top-down modulation of posterior representations.
PMCID: PMC2617739  PMID: 17073983
Brain imaging; dyslexia; learning difficulties; magnetic resonance imaging; phonological processing; reading disorder

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