Overlapping and unique brain responses to cognitive and response inhibition.

Although cognitive inhibition and response inhibition fall under the umbrella term of inhibition, the question remains whether the two aspects of inhibition engage shared or distinct brain regions. The current study is one of the first to examine the neural underpinnings of cognitive inhibition (e.g. the Stroop incongruency effect) and response inhibition (e.g. "no-go" response) within a single task. Adult participants (n = 77) completed an adapted version of the Simon Task in a 3T MRI scanner. The results demonstrated that cognitive and response inhibition recruited a group of overlapping brain regions (inferior frontal cortex, inferior temporal lobe, precentral cortex, parietal cortex). However, a direct comparison of cognitive and response inhibition revealed that the two aspects of inhibition also engaged distinct, task-specific brain regions (voxel-wise FWE corrected p < 0.05). Cognitive inhibition was associated with increases in multiple brain regions within the prefrontal cortex. On the other hand, response inhibition was associated with increases in distinct regions of the prefrontal cortex, right superior parietal cortex, and inferior temporal lobe. Our findings advance the understanding of the brain basis of inhibition by suggesting that cognitive inhibition and response inhibition engage overlapping but distinct brain regions.

Correspondence between cognitive and neural representations for phonology, orthography, and semantics in supramarginal compared to angular gyrus.

The angular and supramarginal gyri (AG and SMG) together constitute the inferior parietal lobule (IPL) and have been associated with cognitive functions that support reading. How those functions are distributed across the AG and SMG is a matter of debate, the resolution of which is hampered by inconsistencies across stereotactic atlases provided by the major brain image analysis software packages. Schematic results from automated meta-analyses suggest primarily semantic (word meaning) processing in the left AG, with more spatial overlap among phonological (auditory word form), orthographic (visual word form), and semantic processing in the left SMG. To systematically test for correspondence between patterns of neural activation and phonological, orthographic, and semantic representations, we re-analyze a functional magnetic resonance imaging data set of participants reading aloud 465 words. Using representational similarity analysis, we test the hypothesis that within cytoarchitecture-defined subregions of the IPL, phonological representations are primarily associated with the SMG, while semantic representations are primarily associated with the AG. To the extent that orthographic representations can be de-correlated from phonological representations, they will be associated with cortex peripheral to the IPL, such as the intraparietal sulcus. Results largely confirmed these hypotheses, with some nuanced exceptions, which we discuss in terms of neurally inspired computational cognitive models of reading that learn mappings among distributed representations for orthography, phonology, and semantics. De-correlating constituent representations making up complex cognitive processes, such as reading, by careful selection of stimuli, representational formats, and analysis techniques, are promising approaches for bringing additional clarity to brain structure-function relationships.

Phonological Preparation in Korean: Phoneme, or Syllable or Another Unit?

With three experiments, the present study investigated the primary phonological preparation (PP) unit in spoken word production in Korean. Adopting the form preparation paradigm, 23 native Korean speakers named pictures in homogenous or heterogenous lists. In homogenous lists, the names of the pictures shared the same initial phoneme (Experiment 1), initial consonant + vowel (i.e., CV) body (Experiment 2), or initial consonant + vowel + consonant (CVC) syllable (Experiment 3); and in heterogenous lists, the names did not share any phonological components systematically. Compared to naming pictures in heterogenous lists, participants' naming speed was significantly faster when the initial body or the initial syllable of target names was shared. However, this form preparation effect was not shown in Experiment 1, when only the initial phoneme was shared. These results suggested that the body serves as the primary PP unit in Korean, that is, native Korean speakers tend to plan spoken words in a body-coda fashion, probably due to a joint contribution from the strong prevalence of the CV structure and early literacy instructional approach.

Prefrontal cortical thickness mediates the association between cortisol reactivity and executive function in childhood.

The impact of stress hormones, such as cortisol, on the brain is proposed to contribute to differences in executive function of school-age children from impoverished backgrounds. However, the association between cortisol reactivity, prefrontal cortex, and executive function is relatively unexplored in young children. The current longitudinal study examined whether 63 children's early preschool-age (3-5 years, Time 1) and concurrent school-age (5-9 years, Time 2) salivary cortisol reactivity were associated with executive function and prefrontal cortical thickness at school-age. Two measures of cortisol reactivity were calculated: area under the curve with respect to ground (AUCg; total cortisol release) and with respect to increase (AUCi; total change in cortisol). Results demonstrated that Time 2 total cortisol release was negatively associated with executive function, Time 1 total cortisol release positively related to right middle frontal cortical thickness, and Time 2 total cortisol change was negatively associated with right inferior frontal cortical thickness. Moreover, greater right middle frontal cortical thickness mediated the association between greater Time 1 total cortisol release and lower executive function. This study provides support for an early adversity framework in which individual differences in executive function in childhood are directly related to the variations of cortisol-release and the effects on the prefrontal cortex thickness.

Interactive activation and mutual constraint satisfaction in perception and cognition.

In a seminal 1977 article, Rumelhart argued that perception required the simultaneous use of multiple sources of information, allowing perceivers to optimally interpret sensory information at many levels of representation in real time as information arrives. Building on Rumelhart's arguments, we present the Interactive Activation hypothesis-the idea that the mechanism used in perception and comprehension to achieve these feats exploits an interactive activation process implemented through the bidirectional propagation of activation among simple processing units. We then examine the interactive activation model of letter and word perception and the TRACE model of speech perception, as early attempts to explore this hypothesis, and review the experimental evidence relevant to their assumptions and predictions. We consider how well these models address the computational challenge posed by the problem of perception, and we consider how consistent they are with evidence from behavioral experiments. We examine empirical and theoretical controversies surrounding the idea of interactive processing, including a controversy that swirls around the relationship between interactive computation and optimal Bayesian inference. Some of the implementation details of early versions of interactive activation models caused deviation from optimality and from aspects of human performance data. More recent versions of these models, however, overcome these deficiencies. Among these is a model called the multinomial interactive activation model, which explicitly links interactive activation and Bayesian computations. We also review evidence from neurophysiological and neuroimaging studies supporting the view that interactive processing is a characteristic of the perceptual processing machinery in the brain. In sum, we argue that a computational analysis, as well as behavioral and neuroscience evidence, all support the Interactive Activation hypothesis. The evidence suggests that contemporary versions of models based on the idea of interactive activation continue to provide a basis for efforts to achieve a fuller understanding of the process of perception.

The neurophysiological bases of EEG and EEG measurement: a review for the rest of us.

A thorough understanding of the EEG signal and its measurement is necessary to produce high quality data and to draw accurate conclusions from those data. However, publications that discuss relevant topics are written for divergent audiences with specific levels of expertise: explanations are either at an abstract level that leaves readers with a fuzzy understanding of the electrophysiology involved, or are at a technical level that requires mastery of the relevant physics to understand. A clear, comprehensive review of the origin and measurement of EEG that bridges these high and low levels of explanation fills a critical gap in the literature and is necessary for promoting better research practices and peer review. The present paper addresses the neurophysiological source of EEG, propagation of the EEG signal, technical aspects of EEG measurement, and implications for interpretation of EEG data.

Measuring working memory is all fun and games: a four-dimensional spatial game predicts cognitive task performance.

We developed a novel four-dimensional spatial task called Shapebuilder and used it to predict performance on a wide variety of cognitive tasks. In six experiments, we illustrate that Shapebuilder: (1) Loads on a common factor with complex working memory (WM) span tasks and that it predicts performance on quantitative reasoning tasks and Ravens Progressive Matrices (Experiment 1), (2) Correlates well with traditional complex WM span tasks (Experiment 2), predicts performance on the conditional go/no go task (Experiment 3) and N-back (Experiment 4), and showed weak or nonsignificant correlations with the Attention Networks Task (Experiment 5), and task switching (Experiment 6). Shapebuilder shows that it exhibits minimal skew and kurtosis, and shows good reliability. We argue that Shapebuilder has many advantages over existing measures of WM, including the fact that it is largely language independent, is not prone to ceiling effects, and take less than 6 min to complete on average.

Using a high-dimensional graph of semantic space to model relationships among words.

The GOLD model (Graph Of Language Distribution) is a network model constructed based on co-occurrence in a large corpus of natural language that may be used to explore what information may be present in a graph-structured model of language, and what information may be extracted through theoretically-driven algorithms as well as standard graph analysis methods. The present study will employ GOLD to examine two types of relationship between words: semantic similarity and associative relatedness. Semantic similarity refers to the degree of overlap in meaning between words, while associative relatedness refers to the degree to which two words occur in the same schematic context. It is expected that a graph structured model of language constructed based on co-occurrence should easily capture associative relatedness, because this type of relationship is thought to be present directly in lexical co-occurrence. However, it is hypothesized that semantic similarity may be extracted from the intersection of the set of first-order connections, because two words that are semantically similar may occupy similar thematic or syntactic roles across contexts and thus would co-occur lexically with the same set of nodes. Two versions the GOLD model that differed in terms of the co-occurence window, bigGOLD at the paragraph level and smallGOLD at the adjacent word level, were directly compared to the performance of a well-established distributional model, Latent Semantic Analysis (LSA). The superior performance of the GOLD models (big and small) suggest that a single acquisition and storage mechanism, namely co-occurrence, can account for associative and conceptual relationships between words and is more psychologically plausible than models using singular value decomposition (SVD).

Age, sex, and verbal abilities affect location of linguistic connectivity in ventral visual pathway.

Previous studies have shown that the strength of connectivity between regions can vary depending upon the cognitive demands of a task. In this study, the location of task-dependent connectivity from the primary visual cortex (V1) was examined in 43 children (ages 9-15) performing visual tasks; connectivity maxima were identified for a visual task requiring a linguistic (orthographic) judgment. Age, sex, and verbal IQ interacted to affect maxima location. Increases in age and verbal IQ produced similar shifts in maxima location; in girls, connectivity maxima shifted primarily laterally within the left temporal lobe, whereas the shift was primarily posterior within occipital cortex among boys. A composite map across all subjects shows an expansion in the area of connectivity with age. Results show that the location of visual/linguistic connectivity varies systematically during development, suggesting that both sex differences and developmental changes in V1 connectivity are related to linguistic function.

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

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

Children with reading disability show brain differences in effective connectivity for visual, but not auditory word comprehension.

BACKGROUND: 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. CONCLUSIONS/SIGNIFICANCE: 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.

Children with reading difficulties show differences in brain regions associated with orthographic processing during spoken language processing.

We explored the neural basis of spoken language deficits in children with reading difficulty, specifically focusing on the role of orthography during spoken language processing. We used functional magnetic resonance imaging (fMRI) to examine differences in brain activation between children with reading difficulties (aged 9-to-15 years) and age-matched children with typical achievement during an auditory rhyming task. Both groups showed activation in bilateral superior temporal gyri (BA 42 and 22), a region associated with phonological processing, with no significant between-group differences. Interestingly, typically achieving children, but not children with reading difficulties, showed activation of left fusiform cortex (BA 37), a region implicated in orthographic processing. Furthermore, this activation was significantly greater for typically achieving children compared to those with reading difficulties. These findings suggest that typical children automatically activate orthographic representations during spoken language processing, while those with reading difficulties do not. Follow-up analyses revealed that the intensity of the activation in the fusiform gyrus was associated with significantly stronger behavioral conflict effects in typically achieving children only (i.e., longer latencies to rhyming pairs with orthographically dissimilar endings than to those with identical orthographic endings; jazz-has vs. cat-hat). Finally, for reading disabled children, a positive correlation between left fusiform activation and nonword reading was observed, such that greater access to orthography was related to decoding ability. Taken together, the results suggest that the integration of orthographic and phonological processing is directly related to reading ability.

Neural correlates of priming effects in children during spoken word processing with orthographic demands.

Priming effects were examined in 40 children (9-15 years old) using functional magnetic resonance imaging (fMRI). An orthographic judgment task required participants to determine if two sequentially presented spoken words had the same spelling for the rime. Four lexical conditions were designed: similar orthography and phonology (O(+)P(+)), similar orthography but different phonology (O(+)P(-)), similar phonology but different orthography (O(-)P(+)), and different orthography and phonology (O(-)P(-)). In left superior temporal gyrus, there was lower activation for targets in O(+)P(+) than for those in O(-)P(-) and higher accuracy was correlated with stronger activation across all lexical conditions. These results provide evidence for phonological priming in children and greater elaboration of phonological representations in higher skill children, respectively. In left fusiform gyrus, there was lower activation for targets in O(+)P(+) and O(+)P(-) than for those in O(-)P(-), suggesting that visual similarity resulted in orthographic priming even with only auditory input. In left middle temporal gyrus, there was lower activation for targets in O(+)P(+) than all other lexical conditions, suggesting that converging orthographic and phonological information resulted in a weaker influence on semantic representations. In addition, higher reading skill was correlated with weaker activation in left middle temporal gyrus across all lexical conditions, suggesting that higher skill children rely to a lesser degree on semantics as a compensatory mechanism. Finally, conflict effects but not priming effects were observed in left inferior frontal gyrus, suggesting that this region is involved in resolving conflicting orthographic and phonological information but not in perceptual priming.

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.

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.

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

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

Neural correlates of orthographic and phonological consistency effects in children.

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 role of discourse context in developing word form representations: a paradoxical relation between reading and learning.

To acquire representations of printed words, children must attend to the written form of a word and link this form with the word's pronunciation. When words are read in context, they may be read with less attention to these features, and this can lead to poorer word form retention. Two experiments with young children (ages 5-8 years) confirmed this hypothesis. In our experiments, children attempted to read words they could not previously read, during a self-teaching period, either in context or in isolation. Later they were tested on how well they learned the words as a function of self-teaching condition (isolation or context). Consistent with previous research, children read more words accurately in context than in isolation during self-teaching; however, children had better retention for words learned in isolation. Furthermore, this benefit from learning in isolation was larger for less skilled readers. This effect of poorer word retention when words are learned in context is paradoxical because context has been shown to facilitate word identification. We discuss factors that may influence this effect of context, especially the role of children's skill level and the demands of learning new word representations at the beginning of reading instruction.

Cross-cultural effect on the brain revisited: universal structures plus writing system variation.

Recognizing printed words requires the mapping of graphic forms, which vary with writing systems, to linguistic forms, which vary with languages. Using a newly developed meta-analytic approach, aggregated Gaussian-estimated sources (AGES; Chein et al. [2002]: Psychol Behav 77:635-639), we examined the neuroimaging results for word reading within and across writing systems and languages. To find commonalities, we compiled 25 studies in English and other Western European languages that use an alphabetic writing system, 9 studies of native Chinese reading, 5 studies of Japanese Kana (syllabic) reading, and 4 studies of Kanji (morpho-syllabic) reading. Using the AGES approach, we created meta-images within each writing system, isolated reliable foci of activation, and compared findings across writing systems and languages. The results suggest that these writing systems utilize a common network of regions in word processing. Writing systems engage largely the same systems in terms of gross cortical regions, but localization within those regions suggests differences across writing systems. In particular, the region known as the visual word form area (VWFA) shows strikingly consistent localization across tasks and across writing systems. This region in the left mid-fusiform gyrus is critical to word recognition across writing systems and languages.