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.     

Developmental changes in activation and effective connectivity in phonological processing

The current study examined developmental changes in activation and effective connectivity among brain regions during a phonological processing task, using fMRI. Participants, ages 9-15, were scanned while performing rhyming judgments on pairs of visually presented words. The orthographic and phonological similarity between words in the pair was independently manipulated, so that rhyming judgment could not be based on orthographic similarity. Our results show a developmental increase in activation in the dorsal part of left inferior frontal gyrus (IFG), accompanied by a decrease in the dorsal part of left superior temporal gyrus (STG). The coupling of dorsal IFG with other selected brain regions involved in the phonological decision increased with age, while the coupling of STG decreased with age. These results suggest that during development there is a shift from reliance on sensory auditory representations to reliance on phonological segmentation and covert articulation for performing rhyming judgment on visually presented words. In addition, we found a developmental increase in activation in left posterior parietal cortex that was not accompanied by a change in its connectivity with the other regions. These results suggest that maturational changes within a cortical region are not necessarily accompanied by an increase in its interactions with other regions and its contribution to the task. Our results are consistent with the idea that there is reduced reliance on primary sensory processes as task-relevant processes mature and become more efficient during development.     

Functional specialization for semantic and phonological processing in the left inferior prefrontal cortex.

Neuroimaging and neuropsychological studies have implicated left inferior prefrontal cortex (LIPC) in both semantic and phonological processing. In this study, functional magnetic resonance imaging was used to examine whether separate LIPC regions participate in each of these types of processing. Performance of a semantic decision task resulted in extensive LIPC activation compared to a perceptual control task. Phonological processing of words and pseudowords in a syllable-counting task resulted in activation of the dorsal aspect of the left inferior frontal gyrus near the inferior frontal sulcus (BA 44/45) compared to a perceptual control task, with greater activation for nonwords compared to words. In a direct comparison of semantic and phonological tasks, semantic processing preferentially activated the ventral aspect of the left inferior frontal gyrus (BA 47/45). A review of the literature demonstrated a similar distinction between left prefrontal regions involved in semantic processing and phonological/lexical processing. The results suggest that a distinct region in the left inferior frontal cortex is involved in semantic processing, whereas other regions may subserve phonological processes engaged during both semantic and phonological tasks.     

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.     

Overlapping neural regions for processing rapid temporal cues in speech and nonspeech signals

Speech perception involves recovering the phonetic form of speech from a dynamic auditory signal containing both time-varying and steady-state cues. We examined the roles of inferior frontal and superior temporal cortex in processing these aspects of auditory speech and nonspeech signals. Event-related functional magnetic resonance imaging was used to record activation in superior temporal gyrus (STG) and inferior frontal gyrus (IFG) while participants discriminated pairs of either speech syllables or nonspeech tones. Speech stimuli differed in either the consonant or the vowel portion of the syllable, whereas the nonspeech signals consisted of sinewave tones differing along either a dynamic or a spectral dimension. Analyses failed to identify regions of activation that clearly contrasted the speech and nonspeech conditions. However, we did identify regions in the posterior portion of left and right STG and left IFG yielding greater activation for both speech and nonspeech conditions that involved rapid temporal discrimination, compared to speech and nonspeech conditions involving spectral discrimination. The results suggest that, when semantic and lexical factors are adequately ruled out, there is significant overlap in the brain regions involved in processing the rapid temporal characteristics of both speech and nonspeech signals.     

Functional MRI study of semantic and phonological language processing in bilingual subjects: preliminary findings

The objective of the study was to explore differences in regional fMRI activation topography and lateralization between semantic and phonological tasks performed in English and Spanish in bilingual individuals. Eight bilingual (primary Spanish and secondary English-speaking) individuals performed fMRI noun-verb association and rhyming tasks in both Spanish and English. Functional dataset analysis within Statistical Parametric Mapping (SPM99) with overlay on T1-weighted anatomic images was performed. Significantly higher laterality indices were noted in the semantic tasks as compared with the phonological tasks in the anterior regions of interest comprising the frontal and superior temporal lobes. A task subtraction analysis demonstrated right hemispheric (inferior frontal gyrus and supramarginal gyrus) foci of significantly increased activation in the combined language phonological tasks compared to the combined language semantic tasks; similarly prominent right hemispheric activation was seen in the English phonological-English semantic subtraction, but the analogous Spanish task subtraction revealed no task-related differences. This divergence in activation topography between semantic and phonological tasks performed in the nonnative language, but not in the primary language, suggests that neural networks utilized for phonological and semantic language processing in the nonnative language may not be as similar as those in the primary language.     

Phonological processing in relation to reading: an fMRI study in deaf readers

Without special education, early deprivation of auditory speech input, hinders the development of phonological representations and may alter the neural mechanisms of reading. By using fMRI during lexical and rhyming decision tasks, we compared in hearing and pre-lingually deaf subjects the neural activity in functional regions of interest (ROIs) engaged in reading. The results show in deaf readers significantly higher activation in the ROIs relevant to the grapho-phonological route, but also in the posterior medial frontal cortex (pMFC) and the right inferior frontal gyrus (IFG). These adjustments may be interpreted within the dual route model of reading as an alternative strategy, which gives priority to rule-based letter-to-sound conversion. Activation in the right IFG would account for compensation mechanisms based on phonological recoding and inner speech while activation in the posterior medial frontal cortex (pMFC) may relate to the cognitive effort called for by the alternative strategy. Our data suggest that the neural mechanisms of reading are shaped by the auditory experience of speech.     

左侧额下回在汉字真假词视觉加工中的不同效应

目的利用事件相关功能磁共振成像(functionalMRI)技术比较中文双字真词与假词的认知加工的脑机制。方法14名大学生在MR扫描过程中执行词汇判断任务,用视觉方式随机呈现120个双字词,其中真、假词各60个。结果真、假词的判断激活了相似的包括双侧额下回,颞中回,梭状回和舌回,双侧缘上回,丘脑等区域。其中,左侧额下回(BA44/45区)在假词条件下的激活强度显著高于真词。结论左额下回可能与语音存贮、行为选择或调控注意有关。     

Cortical activation to auditory mismatch elicited by frequency deviant and complex novel sounds: a PET study

The analysis of auditory deviant events outside the focus of attention is a fundamental capacity of human information processing and has been studied in experiments on Mismatch Negativity (MMN) and the P3a component in evoked potential research. However, generators contributing to these components are still under discussion. Here we assessed cortical blood flow to auditory stimulation in three conditions. Six healthy subjects were presented with standard tones, frequency deviant tones (MMN condition), and complex novel sounds (Novelty condition), while attention was directed to a nondemanding visual task. Analysis of the MMN condition contrasted with thestandard condition revealed blood flow changes in the left and right superior temporal gyrus, right superior temporal sulcus and left inferior frontal gyrus. Complex novel sounds contrasted with the standard condition activated the left superior temporal gyrus and the left inferior and middle frontal gyrus. A small subcortical activation emerged in the left parahippocampal gyrus and an extended activation was found covering the right superior temporal gyrus. Novel sounds activated the right inferior frontal gyrus when controlling for deviance probability. In contrast to previous studies our results indicate a left hemisphere contribution to a frontotemporal network of auditory deviance processing. Our results provide further evidence for a contribution of the frontal cortex to the processing of auditory deviance outside the focus of directed attention.     

Auditory Selective Attention: An fMRI Investigation

In the present experiment, 25 adult subjects discriminated speech tokens ([ba]/[da]) or made pitch judgments on tone stimuli (rising/falling) under both binaural and dichotic listening conditions. We observed that when listeners performed tasks under the dichotic conditions, during which greater demands are made on auditory selective attention, activation within the posterior (parietal) attention system and at primary processing sites in the superior temporal and inferior frontal regions was increased. The cingulate gyrus within the anterior attention system was not influenced by this manipulation. Hemispheric differences between speech and nonspeech tasks were also observed, both at Broca's Area within the inferior frontal gyrus and in the middle temporal gyrus.     

Neuroimaging reveals automatic speech coding during perception of written word meaning

The extent to which visual word perception engages speech codes (i.e., phonological recoding) remains a crucial question in understanding mechanisms of reading. In this study, we used functional magnetic resonance imaging techniques combined with behavioral response measures to examine neural responses to focused versus incidental phonological and semantic processing of written words. Three groups of subjects made simple button-pressing responses in either phonologically (rhyming-judgment) or semantically (category-judgment) focused tasks or both tasks with identical sets of visual stimuli. In the phonological tasks, subjects were given both words and pseudowords separated in different scan runs. The baseline task required feature search of scrambled letter strings created from the stimuli for the experimental conditions. The results showed that cortical regions associated with both semantic and phonological processes were strongly activated when the task required active processing of word meaning. However, when subjects were actively processing the speech sounds of the same set of written words, brain areas typically engaged in semantic processing became silent. In addition, subjects who performed both the rhyming and the semantic tasks showed diverse and significant bilateral activation in the prefrontal, temporal, and other brain regions. Taken together, the pattern of brain activity provides evidence of a neural basis supporting the theory that in normal word reading, phonological recoding is automatic and facilitates semantic processing of written words, while rapid comprehension of word meaning requires devoted attention. These results also raise questions about including multiple cognitive tasks in the same neuroimaging sessions.     

Working memory in another dimension: functional imaging of human olfactory working memory

The majority of working memory research has been carried out within the visual and auditory modalities, leaving it unclear how other modalities would map onto currently proposed working memory models. In this study we examined the previously uninvestigated area of olfactory working memory. Our aim was to investigate if olfactory working memory would engage prefrontal regions known to be involved in working memory for other sensory modalities. Using positron emission tomography we measured cerebral blood flow changes in 12 volunteers during an olfactory working memory task and a comparison visual working memory task. Our findings indicate that both olfactory and face working memory engaged dorsolateral and ventrolateral frontal cortex when the task requirements were matched; a conjunction analysis indicated overlap in the distribution of activity in the two tasks. Similarities and differences in activity were noted in parietal lobe regions, with both tasks engaging inferior areas of 40/7, but only visual working memory showing increased activity within left superior parietal cortex. The findings support the idea that working memory processes engage frontal cortical areas independent of the modality of input, but do not rule out the possibility of modality-specific neural populations within dorsolateral or ventrolateral cortex.     

Time-dependent changes in learning audiovisual associations: a single-trial fMRI study

Functional imaging studies of learning and memory have primarily focused on stimulus material presented within a single modality (see review by Gabrieli, 1998, Annu. Rev. Psychol. 49: 87-115). In the present study we investigated mechanisms for learning material presented in visual and auditory modalities, using single-trial functional magnetic resonance imaging. We evaluated time-dependent learning effects under two conditions involving presentation of consistent (repeatedly paired in the same combination) or inconsistent (items presented randomly paired) pairs. We also evaluated time-dependent changes for bimodal (auditory and visual) presentations relative to a condition in which auditory stimuli were repeatedly presented alone. Using a time by condition analysis to compare neural responses to consistent versus inconsistent audiovisual pairs, we found significant time-dependent learning effects in medial parietal and right dorsolateral prefrontal cortices. In contrast, time-dependent effects were seen in left angular gyrus, bilateral anterior cingulate gyrus, and occipital areas bilaterally. A comparison of paired (bimodal) versus unpaired (unimodal) conditions was associated with time-dependent changes in posterior hippocampal and superior frontal regions for both consistent and inconsistent pairs. The results provide evidence that associative learning for stimuli presented in different sensory modalities is supported by neural mechanisms similar to those described for other kinds of memory processes. The involvement of posterior hippocampus and superior frontal gyrus in bimodal learning for both consistent and inconsistent pairs supports a putative function for these regions in associative learning independent of sensory modality.     

Orthographic and phonological processing of Chinese characters: an fMRI study

The present study used functional magnetic resonance imaging (fMRI) to investigate the neural mechanisms underlying the orthographic and phonological processing of Chinese characters. Four tasks were devised, including one homophone judgment and three physical judgments of characters, pseudo-characters, and Korean-like nonsense figures. While the left occipitotemporal region, left dorsal processing stream, and right middle frontal gyrus constitute a network for orthographic processing, the left premotor gyrus, left middle/inferior frontal gyrus, supplementary motor area (SMA), and the left temporoparietal region work in concert for phonological processing. The ventral part of the left inferior frontal cortex responds specifically to the character stimuli, suggesting a general lexical processing role for this region for linguistic material. The stronger activation of the dorsal visual stream by Chinese homophone judgment pinpoints a tight coupling between phonological representation of Chinese characters and corresponding orthographic percepts. The concomitant engagement of sets of regions for different levels of Chinese orthographic and phonological processing is consistent with the notion of distributed parallel processing.     

Imaging phonology without print: assessing the neural correlates of phonemic awareness using fMRI

Acquisition of phonological processing skills, such as the ability to segment words into corresponding speech sounds, is critical to the development of efficient reading. Prior neuroimaging studies of phonological processing have often relied on auditory stimuli or print-mediated tasks that may be problematic for various theoretical and empirical reasons. For the current study, we developed a task to evaluate phonological processing that used visual stimuli but did not require interpretation of orthographic forms. This task requires the subject to retrieve the names of objects and to compare their first sounds; then, the subject must indicate if the initial sounds of the names of the pictures are the same. The phonological analysis task was compared to both a baseline matching task and a more complex control condition in which the participants evaluated two different pictures and indicated whether they represented the same object. The complex picture-matching condition controls for the visual complexity of the stimuli but does not require phonological analysis of the names of the objects. While both frontal and ventral posterior areas were activated in response to phonological analysis of the names of pictures, only inferior and superior frontal gyrus exhibited differential sensitivity to the phonological comparison task as compared to the complex picture-matching control task. These findings suggest that phonological processing that is not mediated by print relies primarily on frontal language processing areas among skilled readers.     

Changing channels: an fMRI study of aging and cross-modal attention shifts

Age-related deficits in visual selective attention suggest that the efficiency of inhibitory processes is particularly affected by aging. To investigate whether processing inefficiencies observed in visual attention are similar in auditory attention and when shifting attention across modalities, we conducted an fMRI study with healthy young and older adults using a task that required sustained auditory and visual selective attention and cross-modal attention shifts. Older adults in this study performed as well as the younger adults, but showed age-related differences in BOLD responses. The most striking of these differences were bilateral frontal and parietal regions of significantly increased activation in older adults during both focused and shifting attention. Our data suggest that this increased activation did not reflect new recruitment, but reliance on brain regions typically used by younger adults when task demands are greater. Older adults' activation patterns suggested that even during focused attention conditions they were "shifting" attention to stimuli in the unattended modality. Increased activation during processing of both task-relevant and task-irrelevant information implies age-related loss of processing selectivity. These patterns may reflect both task-specific compensatory neural recruitment and degradation of sensory inhibition.     

Temporal and topographical characterization of language cortices using intraoperative optical intrinsic signals

We used intraoperative optical imaging of intrinsic signals (iOIS) and electrocortical stimulation mapping (ESM) to compare functionally active brain regions in 10 awake patients undergoing neurosurgical resection. Patients performed two to four tasks, including visual and auditory naming, word discrimination, and/or orofacial movements. All iOIS maps included areas identified by ESM mapping. However, iOIS also revealed topographical specificity dependent on language task. In Broca's area, naming paradigms activated both anterior and posterior inferior frontal gyrus (IFG), while the word discrimination paradigm activated only posterior IFG. In Wernicke's area, object naming produced activations localizing over the inferior and anterior/posterior regions, while the word discrimination task activated superior and anterior cortices. These results may suggest more posterior phonological activation and more anterior semantic activations in Broca's area, and more anterior/superior phonological activation and more posterior/inferior semantic activations in Wernicke's area. Although similar response onset was observed in Broca's and Wernicke's areas, temporal differences were revealed during block paradigm (20-s) activations. In Broca's area, block paradigms yielded a boxcar temporal activation profile (in all tasks) that resembled response profiles observed in motor cortex (with orofacial movements). In contrast, activations in Wernicke's area responded with a more dynamic profile (including early and late peaks) which varied with paradigm performance. Wernicke's area profiles were very similar to response profiles observed in sensory and visual cortex. The differing temporal patterns may therefore reflect unique processing performed by receptive (Wernicke's) and productive (Broca's) language centers. This study is consistent with task-specific semantic and phonologic regions within Broca's and Wernicke's areas and also is the first report of response profile differences dependent on cortical region and language task.     

A meta-analysis of fMRI studies on Chinese orthographic, phonological, and semantic processing

A growing body of neuroimaging evidence has shown that Chinese character processing recruits differential activation from alphabetic languages due to its unique linguistic features. As more investigations on Chinese character processing have recently become available, we applied a meta-analytic approach to summarize previous findings and examined the neural networks for orthographic, phonological, and semantic processing of Chinese characters independently. The activation likelihood estimation (ALE) method was used to analyze eight studies in the orthographic task category, eleven in the phonological and fifteen in the semantic task categories. Converging activation among three language-processing components was found in the left middle frontal gyrus, the left superior parietal lobule and the left mid-fusiform gyrus, suggesting a common sub-network underlying the character recognition process regardless of the task nature. With increasing task demands, the left inferior parietal lobule and the right superior temporal gyrus were specialized for phonological processing, while the left middle temporal gyrus was involved in semantic processing. Functional dissociation was identified in the left inferior frontal gyrus, with the posterior dorsal part for phonological processing and the anterior ventral part for semantic processing. Moreover, bilateral involvement of the ventral occipito-temporal regions was found for both phonological and semantic processing. The results provide better understanding of the neural networks underlying Chinese orthographic, phonological, and semantic processing, and consolidate the findings of additional recruitment of the left middle frontal gyrus and the right fusiform gyrus for Chinese character processing as compared with the universal language network that has been based on alphabetic languages.     

Functional neuroanatomy of auditory mismatch processing: an event-related fMRI study of duration-deviant oddballs

This study was designed to identify the neural networks underlying automatic auditory deviance detection in 10 healthy subjects using functional magnetic resonance imaging. We measured blood oxygenation level-dependent contrasts derived from the comparison of blocks of stimuli presented as a series of standard tones (50 ms duration) alone versus blocks that contained rare duration-deviant tones (100 ms) that were interspersed among a series of frequent standard tones while subjects were watching a silent movie. Possible effects of scanner noise were assessed by a "no tone" condition. In line with previous positron emission tomography and EEG source modeling studies, we found temporal lobe and prefrontal cortical activation that was associated with auditory duration mismatch processing. Data were also analyzed employing an event-related hemodynamic response model, which confirmed activation in response to duration-deviant tones bilaterally in the superior temporal gyrus and prefrontally in the right inferior and middle frontal gyri. In line with previous electrophysiological reports, mismatch activation of these brain regions was significantly correlated with age. These findings suggest a close relationship of the event-related hemodynamic response pattern with the corresponding electrophysiological activity underlying the event-related "mismatch negativity" potential, a putative measure of auditory sensory memory.     

Functional anatomy of pitch memory--an fMRI study with sparse temporal sampling

Auditory functional magnetic resonance imaging tasks are challenging since the MR scanner noise can interfere with the auditory stimulation. To avoid this interference a sparse temporal sampling method with a long repetition time (TR = 17 s) was used to explore the functional anatomy of pitch memory. Eighteen right-handed subjects listened to a sequence of sine-wave tones (4.6 s total duration) and were asked to make a decision (depending on a visual prompt) whether the last or second to last tone was the same or different as the first tone. An alternating button press condition served as a control. Sets of 24 axial slices were acquired with a variable delay time (between 0 and 6 s) between the end of the auditory stimulation and the MR acquisition. Individual imaging time points were combined into three clusters (0-2, 3-4, and 5-6 s after the end of the auditory stimulation) for the analysis. The analysis showed a dynamic activation pattern over time which involved the superior temporal gyrus, supramarginal gyrus, posterior dorsolateral frontal regions, superior parietal regions, and dorsolateral cerebellar regions bilaterally as well as the left inferior frontal gyrus. By regressing the performance score in the pitch memory task with task-related MR signal changes, the supramarginal gyrus (left>right) and the dorsolateral cerebellum (lobules V and VI, left>right) were significantly correlated with good task performance. The SMG and the dorsolateral cerebellum may play a critical role in short-term storage of pitch information and the continuous pitch discrimination necessary for performing this pitch memory task.