Douglas D. Burman

Functional Anatomy of Intra- and Cross-Modal Lexical Tasks

Functional magnetic resonance imaging (fMRI) was used to examine lexical processing in normal adults (20-35 years). Two tasks required only intramodal processing (spelling judgments with visual input and rhyming judgments with auditory input) and two tasks required cross-modal processing between phonologic and orthographic representations (spelling judgments with auditory input and rhyming judgments with visual input). Each task led to greater activation in the unimodal association area concordant with the modality of input, namely fusiform gyrus (BA 19, 37) for written words and superior temporal gyrus (BA 22, 42) for spoken words. Cross-modal tasks generated greater activation in posterior heteromodal regions including the supramarginal and angular gyri (BA 40, 39). Cross-modal tasks generated additional activation in unimodal areas representing the target of conversion, superior temporal gyrus for visual rhyming and fusiform gyrus for auditory spelling. Our findings suggest that the fusiform gyrus processes orthographic word forms, the superior temporal gyrus processes phonologic word forms, and posterior heteromodal regions are involved in the conversion between orthography and phonology.

Neural development of selective attention and response inhibition

Brain activation differences between 12 children (9- to 12-year-olds) and 12 adults (20- to 30-year-olds) were examined on two cognitive tasks during functional magnetic resonance imaging (fMRI). Spatial selective attention was measured with the visual search for a conjunction target (red triangle) in a field of distracters and response inhibition was measured with a go no-go task. There were small developmental differences in the selective attention task, with children showing greater activation than adults in the anterior cingulate and thalamus. There were large developmental differences in the response inhibition task, with children showing greater activation than adults in a fronto-striatal network including middle cingulate, medial frontal gyrus, medial aspects of bilateral superior frontal gyrus, and the caudate nucleus on the left. Children also showed greater bilateral activation for the response inhibition task in posterior cingulate, thalamus and the hippocampo-amygdaloid region. The extensive developmental differences on the response inhibition task are consistent with the prolonged maturation of the fronto-striatal network.

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). Hum. Brain Mapping 16:251–261, 2002.

Development of Brain Mechanisms for Processing Orthographic and Phonologic Representations

Developmental differences in the neurocognitive networks for lexical processing were examined in 15 adults and 15 children (9-to 12-year-olds) using functional magnetic resonance imaging (fMRI). The lexical tasks involved spelling and rhyming judgments in either the visual or auditory modality. These lexical tasks were compared with nonlinguistic control tasks involving judgments of line patterns or tone sequences. The first main finding was that adults showed greater activation than children during the cross-modal lexical tasks in a region proposed to be involved in mapping between orthographic and phonologic representations. The visual rhyming task, which required conversion from orthography to phonology, produced greater activation for adults in the angular gyrus. The auditory spelling task, which required the conversion from phonology to orthography, also produced greater activation for adults in the angular gyrus. The greater activation for adults suggests they may have a more elaborated posterior heteromodal system for mapping between representational systems. The second main finding was that adults showed greater activation than children during the intra-modal lexical tasks in the angular gyrus. The visual spelling and auditory rhyming did not require conversion between orthography and phonology for correct performance but the adults showed greater activation in a system implicated for this mapping. The greater activation for adults suggests that they have more interactive convergence between representational systems during lexical processing.

Shifts of Effective Connectivity within a Language Network during Rhyming and Spelling

We used functional magnetic resonance imaging to examine task-specific modulations of effective connectivity within a left-hemisphere language network during spelling and rhyming judgments on visually presented words. We identified sites showing task-specific activations for rhyming in the lateral temporal cortex (LTC) and for spelling in the intraparietal sulcus (IPS). The inferior frontal gyrus (IFG) and fusiform gyrus were engaged by both tasks. Dynamic causal modeling showed that each task preferentially strengthened modulatory influences converging on its task-specific site (LTC for rhyming, IPS for spelling). These remarkably selective and symmetrical findings demonstrate that the nature of the behavioral task dynamically shifts the locus of integration (or convergence) to the network component specialized for that task. Furthermore, they suggest that the role of the task-selective areas is to provide a differential synthesis of incoming information rather than providing differential control signals influencing the activity of other network components. Our findings also showed that switching tasks led to changes in the target area influenced by the IFG, suggesting that the IFG may play a pivotal role in setting the cognitive context for each task. We propose that task-dependent shifts in effective connectivity are likely to be mediated through top-down modulations from the IFG to the task-selective regions in a way that differentially enhances their sensitivity to incoming word-form information.

Sex Differences in Neural Processing of Language Among Children

Why females generally perform better on language tasks than males is unknown. Sex differences were here identified in children (ages 9-15) across two linguistic tasks for words presented in two modalities. Bilateral activation in the inferior frontal and superior temporal gyri and activation in the left fusiform gyrus of girls was greater than in boys. Activation in the left inferior frontal and fusiform regions of girls was also correlated with linguistic accuracy irregardless of stimulus modality, whereas correlation with performance accuracy in boys depended on the modality of word presentation (either in visual or auditory association cortex). This pattern suggests that girls rely on a supramodal language network, whereas boys process visual and auditory words differently. Activation in the left fusiform region was additionally correlated with performance on standardized language tests in which girls performed better, additional evidence of its role in early sex differences for language.

Relation between brain activation and lexical performance

Functional magnetic resonance imaging (fMRI) was used to determine whether performance on lexical tasks was correlated with cerebral activation patterns. We found that such relationships did exist and that their anatomical distribution reflected the neurocognitive processing routes required by the task. Better performance on intramodal tasks (determining if visual words were spelled the same or if auditory words rhymed) was correlated with more activation in unimodal regions corresponding to the modality of sensory input, namely the fusiform gyrus (BA 37) for written words and the superior temporal gyrus (BA 22) for spoken words. Better performance in tasks requiring cross‐modal conversions (determining if auditory words were spelled the same or if visual words rhymed), on the other hand, was correlated with more activation in posterior heteromodal regions, including the supramarginal gyrus (BA 40) and the angular gyrus (BA 39). Better performance in these cross‐modal tasks was also correlated with greater activation in unimodal regions corresponding to the target modality of the conversion process (i.e., fusiform gyrus for auditory spelling and superior temporal gyrus for visual rhyming). In contrast, performance on the auditory spelling task was inversely correlated with activation in the superior temporal gyrus possibly reflecting a greater emphasis on the properties of the perceptual input rather than on the relevant transmodal conversions. Hum. Brain Mapping 19:155–169, 2003.

The Development of Specialized Brain Systems in Reading and Oral-Language

Functional magnetic resonance imaging (fMRI) was used to examine differences between children (9–12 years) and adults (21–31 years) in the distribution of brain activation during word processing. Orthographic, phonologic, semantic and syntactic tasks were used in both the auditory and visual modalities. Our two principal results were consistent with the hypothesis that development is characterized by increasing specialization. Our first analysis compared activation in children versus adults separately for each modality. Adults showed more activation than children in the unimodal visual areas of middle temporal gyrus and fusiform gyrus for processing written word forms and in the unimodal auditory areas of superior temporal gyrus for processing spoken word forms. Children showed more activation than adults for written word forms in posterior heteromodal regions (Wernickes area), presumably for the integration of orthographic and phonologic word forms. Our second analysis compared activation in the visual versus auditory modality separately for children and adults. Children showed primarily overlap of activation in brain regions for the visual and auditory tasks. Adults showed selective activation in the unimodal auditory areas of superior temporal gyrus when processing spoken word forms and selective activation in the unimodal visual areas of middle temporal gyrus and fusiform gyrus when processing written word forms.

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.

Developmental and Skill Effects on the Neural Correlates of Semantic Processing to Visually Presented Words

Functional magnetic resonance imaging (fMRI) was used to explore the neural correlates of semantic judgments to visual words in a group of 9‐ to 15‐year‐old children. Subjects were asked to indicate if word pairs were related in meaning. Consistent with previous findings in adults, children showed activation in bilateral inferior frontal gyri (Brodmann area [BA] 47, 45) and left middle temporal gyrus (BA 21). Words with strong semantic association elicited significantly greater activation in bilateral inferior parietal lobules (BA 40), suggesting stronger integration of highly related semantic features. By contrast, words with weak semantic association elicited greater activation in left inferior frontal gyrus (BA 45) and middle temporal gyrus (BA 21), suggesting more difficult feature search and more extensive access to semantic representations. We also examined whether age and skill explained unique variance in the patterns of activation. Increasing age was correlated with greater activation in left middle temporal gyrus (BA 21) and inferior parietal lobule (BA 40), suggesting that older children have more elaborated semantic representations and more complete semantic integration processes, respectively. Decreasing age was correlated with activation in right superior temporal gyrus (BA 22) and decreasing accuracy was correlated with activation in right middle temporal gyrus (BA 21), suggesting the engagement of ancillary systems in the right hemisphere for younger and lower‐skill children. Hum. Brain Mapping 2006.