Left hemisphere superiority for pronounceable nonwords,but not for unpronounceable letter strings

Right-handed adults were asked to identify bilaterally presented linguistic stimuli under three experimental conditions. In Condition A, stimuli were three-letter pronounceable nonwords (such as TUP), and subjects were asked to report them by naming them. In Condition B, stimuli were three-letter pronounceable nonwords, and subjects were asked to report them as strings of letters. In Condition C, stimuli were more or less unpronounceable letter strings (such as UTP) created by rearranging the letters of pronounceable nonwords, and subjects reported them as strings of letters. Pronounceable nonwords were found to be better identified from the right visual hemifield irrespective of the way in which they were reported. Unpronounceable letter strings did not produce any visual hemifield difference. Nonwords are of interest because they can be seen as potential words that lack both specific semantic properties and entries in the subject's internal lexicon. The results of the experiment are consistent with the view that both the left and right cerebral hemispheres are able to identify letters but the left hemisphere is more sensitive to the pronounceability of the nonwords. This may happen either because the left hemisphere can make better use of resemblances to real words or because it has access to spelling to sound correspondence rules.     

Auditory language comprehension: an event-related fMRI study on the processing of syntactic and lexical information

The functional specificity of different brain regions recruited in auditory language processing was investigated by means of event-related functional magnetic resonance imaging (fMRI) while subjects listened to speech input varying in the presence or absence of semantic and syntactic information. There were two sentence conditions containing syntactic structure, i.e., normal speech (consisting of function and content words), syntactic speech (consisting of function words and pseudowords), and two word-list conditions, i.e., real words and pseudowords. The processing of auditory language, in general, correlates with significant activation in the primary auditory cortices and in adjacent compartments of the superior temporal gyrus bilaterally. Processing of normal speech appeared to have a special status, as no frontal activation was observed in this case but was seen in the other three conditions. This difference may point toward a certain automaticity of the linguistic processes used during normal speech comprehension. When considering the three other conditions, we found that these were correlated with activation in both left and right frontal cortices. An increase of activation in the planum polare bilaterally and in the deep portion of the left frontal operculum was found exclusively when syntactic processes were in focus. Thus, the present data may be taken to suggest an involvement of the left frontal and bilateral temporal cortex when processing syntactic information during comprehension.     

Functional connectivity in an fMRI study of semantic and phonological processes and the effect of L-Dopa

We describe an fMRI experiment examining the functional connectivity (FC) between regions of the brain associated with semantic and phonological processing. We wished to explore whether L-Dopa administration affects the interaction between language network components in semantic and phonological categorization tasks, as revealed by FC. We hypothesized that L-Dopa would decrease FC due to restriction of the semantic network. During two test sessions (placebo and L-Dopa) each participant performed two fMRI runs, involving phonological and semantic processing. A number of brain regions commonly activated by the two tasks were chosen as regions if interest: left inferior frontal, left posterior temporal and left fusiform gyri, and left parietal cortex. FC was calculated and further analyzed for effects of either the drug or task. No main effect for drug was found. A significant main effect for task was found, with a greater average correlation for the phonological task than for the semantic task. These findings suggest that language areas are activated in a more synchronous manner for phonological than for semantic tasks. This may relate to the fact that phonological processes are mediated to a greater extent within language areas, whereas semantic tasks likely require greater interaction outside of the language areas. Alternatively, this may be due to differences in the attentional requirements of the two tasks.     

Using fMRI to study recovery from acquired dysphasia

We have used functional magnetic resonance imaging (fMRI) to characterize brain activations associated with two distinct language tasks performed by a 28-year-old woman after partial recovery from dysphasia due to a left frontal hemispheric ischemic stroke. MRI showed that her ischemic lesion extended posteriorly from the left inferior frontal to the perisylvian cortex. fMRI scans of both language tasks revealed substantial differences in activation pattern relative to controls. The nature of this difference was task-specific. During performance of a verbal semantic decision task, the patient, in contrast to controls, activated a network of brain areas that excluded the inferior frontal gyrus (in either hemisphere). A second task involving rhyme judgment was designed to place a heavier cognitive load on language production processes and activated the left inferior frontal gyrus (Broca's area) strongly in normal controls. During this task, the most prominent frontal activation in the patient occurred in the right homologue of Broca's area. Subsequent analysis of this data by methods able to deal with responses of changing amplitude revealed additional, less sustained recruitment by the patient of cortex adjacent to the infarct in the region inferior to Broca's area during rhyming. These results suggest that in addition to changes in cognitive strategy, recovery from dysphasia could be mediated by both the preservation of neuronal networks in and around the infarct and the use of homologous regions in the contralateral hemisphere.     

Auditory language comprehension: an event-related fMRI study on the processing of syntactic and lexical information

The functional specificity of different brain areas recruited in auditory language processing was investigated by means of event-related functional magnetic resonance imaging (fMRI) while subjects listened to speech input varying in the presence or absence of semantic and syntactic information. There were two sentence conditions containing syntactic structure, i.e., normal speech (consisting of function and content words), syntactic speech (consisting of function words and pseudowords), and two word-list conditions, i.e., real words and pseudowords. The processing of auditory language, in general, correlates with significant activation in the primary auditory cortices and in adjacent compartments of the superior temporal gyrus bilaterally. Processing of normal speech appeared to have a special status, as no frontal activation was observed in this case but was seen in the three other conditions. This difference may point toward a certain automaticity of the linguistic processes used during normal speech comprehension. When considering the three other conditions, we found that these were correlated with activation in both left and right frontal cortices. An increase of activation in the planum polare bilaterally and in the deep portion of the left frontal operculum was found exclusively when syntactic processes were in focus. Thus, the present data may be taken to suggest an involvement of the left frontal and bilateral temporal cortex when processing syntactic information during comprehension.     

Oral report of words and word approximations presented to the left or right visual field

Subjects reported letter strings forming words, pronounceable high approximations to words, and unpronounceable low approximations to words presented tachistoscopically to the left or right visual field (LVF, RVF). (a) For number of strings totally correct, the same RVF superiority was obtained with high approximations as with words, the field difference with low approximations being negligible. (b) In contrast, for letter scores from partially correct strings, RVF superiority did not vary with string type. Finding (a) is interpreted to indicate that the left hemisphere is differentially specialized for processing words as units and that requiring oral report makes pronounceable strings processable as word-like units. Finding (b) suggests that the left hemisphere is not specialized for processing subword fragments.     

Cerebral organization of oral and signed language responses: case study evidence from amytal and cortical stimulation studies

A normally hearing left-handed patient familiar with American Sign Language (ASL) was assessed under sodium amytal conditions and with left cortical stimulation in both oral speech and signed English. Lateralization was mixed but complementary in each language mode: the right hemisphere perfusion severely disrupted motoric aspects of both types of language expression, the left hemisphere perfusion specifically disrupted features of grammatical and semantic usage in each mode of expression. Both semantic and syntactic aspects of oral and signed responses were altered during left posterior temporal-parietal stimulation. Findings are discussed in terms of the neurological organization of ASL and linguistic organization in cases of early left hemisphere damage.     

Reorganization of the cerebro-cerebellar network of language production in patients with congenital left-hemispheric brain lesions

Patients with congenital lesions of the left cerebral hemisphere may reorganize language functions into the right hemisphere. In these patients, language production is represented homotopically to the left-hemispheric language areas. We studied cerebellar activation in five patients with congenital lesions of the left cerebral hemisphere to assess if the language network is reorganized completely in these patients, i.e. including also cerebellar language functions. As compared to a group of controls matched for age, sex, and verbal IQ, the patients recruited an area not in the right but in the left cerebellar hemisphere. The extent of laterality of the cerebellar activation correlated significantly with the laterality of the frontal activation. We suggest that the developing brain reacts to early focal lesions in the left hemisphere with a mirror-image organization of the entire cerebro-cerebellar network engaged in speech production.     

Effective brain connectivity in children with reading difficulties during phonological processing

Using Dynamic Causal Modeling (DCM) and functional magnetic resonance imaging (fMRI), we examined effective connectivity between three left hemisphere brain regions (inferior frontal gyrus, inferior parietal lobule, fusiform gyrus) and bilateral medial frontal gyrus in 12 children with reading difficulties (M age=12.4, range: 8.11-14.10) and 12 control children (M age=12.3, range: 8.9-14.11) during rhyming judgments to visually presented words. More difficult conflicting trials either had similar orthography but different phonology (e.g. pint-mint) or similar phonology but different orthography (e.g. jazz-has). Easier non-conflicting trials had similar orthography and phonology (e.g. dime-lime) or different orthography and phonology (e.g. staff-gain). The modulatory effect from left fusiform gyrus to left inferior parietal lobule was stronger in controls than in children with reading difficulties only for conflicting trials. Modulatory effects from left fusiform gyrus and left inferior parietal lobule to left inferior frontal gyrus were stronger for conflicting trials than for non-conflicting trials only in control children but not in children with reading difficulties. Modulatory effects from left inferior frontal gyrus to inferior parietal lobule, from medial frontal gyrus to left inferior parietal lobule, and from left inferior parietal lobule to medial frontal gyrus were positively correlated with reading skill only in control children. These findings suggest that children with reading difficulties have deficits in integrating orthography and phonology utilizing left inferior parietal lobule, and in engaging phonological rehearsal/segmentation utilizing left inferior frontal gyrus possibly through the indirect pathway connecting posterior to anterior language processing regions, especially when the orthographic and phonological information is conflicting.     

Neural dichotomy of word concreteness: a view from functional neuroimaging

Our perception about the representation and processing of concrete and abstract concepts is based on the fact that concrete words are highly imagined and remembered faster than abstract words. In order to explain the processing differences between abstract and concrete concepts, various theories have been proposed, yet there is no unanimous consensus about its neural implication. The present study investigated the processing of concrete and abstract words during an orthography judgment task (implicit semantic processing) using functional magnetic resonance imaging to validate the involvement of the neural regions. Relative to non-words, both abstract and concrete words show activation in the regions of bilateral hemisphere previously associated with semantic processing. The common areas (conjunction analyses) observed for abstract and concrete words are bilateral inferior frontal gyrus (BA 44/45), left superior parietal (BA 7), left fusiform gyrus and bilateral middle occipital. The additional areas for abstract words were noticed in bilateral superior temporal and bilateral middle temporal region, whereas no distinct region was noticed for concrete words. This suggests that words with abstract concepts recruit additional language regions in the brain.     

Neuroanatomical correlates of oral reading in acute left hemispheric stroke

Oral reading is a complex skill involving the interaction of orthographic, phonological, and semantic processes. Functional imaging studies with nonimpaired adult readers have identified a widely distributed network of frontal, inferior parietal, posterior temporal, and occipital brain regions involved in the task. However, while functional imaging can identify cortical regions engaged in the process under examination, it cannot identify those brain regions essential for the task. The current study aimed to identify those neuroanatomical regions critical for successful oral reading by examining the relationship between word and nonword oral reading deficits and areas of tissue dysfunction in acute stroke. We evaluated 91 patients with left hemisphere ischemic stroke with a test of oral word and nonword reading, and magnetic resonance diffusion-weighted and perfusion-weighted imaging, within 24-48 h of stroke onset. A voxel-wise statistical map showed that impairments in word and nonword reading were associated with a distributed network of brain regions, including the inferior and middle frontal gyri, the middle temporal gyrus, the supramarginal and angular gyri, and the middle occipital gyrus. In addition, lesions associated with word deficits were found to be distributed more frontally, while nonword deficits were associated with lesions distributed more posteriorly.     

左侧额下回在句子语义整合加工中的作用

语义整合帮助人们在阅读理解中将小信息块整合成一个完整、连贯的句子意义表达, 是阅读理解中非常重要的认知过程。通过对比40多篇句子加工相关的脑机制研究, 发现左侧额下回在fMRI研究中是参与语义整合加工激活概率最高的区域, 而颞叶及后部是MEG研究中激活概率最高的区域。另外, 左侧额下回是如何参与语义整合、它在内隐和外显语义整合中的机制是否相同、以及这种整合加工与一般的控制性加工、词汇启动的关系都是研究者广泛关注的问题。本文对上述问题进行了详细的综述和讨论。     

Fluent versus nonfluent primary progressive aphasia: a comparison of clinical and functional neuroimaging features

To better characterize fluent and nonfluent variants of primary progressive aphasia (PPA). Although investigators have recognized both fluent and nonfluent patients with PPA, the clinical and neuroimaging features of these variants have not been fully defined. We present clinical and neuropsychological data on 47 PPA patients comparing the fluent (n=21) and nonfluent (n=26) subjects. We further compared language features with PET/SPECT data available on 39 of these patients. Compared to the nonfluent PPA patients, those with fluent PPA had greater impairment of confrontational naming and loss of single word comprehension. They also exhibited semantic paraphasic errors and loss of single word comprehension. Patients with nonfluent PPA were more likely to be female, were more often dysarthric, and exhibited phonological speech errors in the absence of semantic errors. No significant differences were seen with regard to left hemisphere abnormalities, suggesting that both variants result from mechanisms that overlap frontal, temporal, and parietal regions. Of the language measures, only semantic paraphasias were strongly localized, in this case to the left temporal lobe. Fluent and nonfluent forms of PPA are clinically distinguishable by letter fluency, single word comprehension, object naming, and types of paraphasic errors. Nevertheless, there is a large amount of overlap between dysfunctional anatomic regions associated with these syndromes.     

The functional anatomy of word comprehension and production

This review describes the functional anatomy of word comprehension and production. Data from functional neuroimaging studies of normal subjects are used to determine the distributed set of brain regions that are engaged during particular language tasks and data from studies of patients with neurological damage are used to determine which of these regions are necessary for task performance. This combination of techniques indicates that the left inferior temporal and left posterior inferior parietal cortices are required for accessing semantic knowledge; the left posterior basal temporal lobe and the left frontal operculum are required for translating semantics into phonological output and the left anterior inferior parietal cortex is required for translating orthography to phonology. Further studies are required to establish the specific functions of the different regions and how these functions interact to provide our sophisticated language system.     

A fMRI study of word retrieval in aphasia

The neural mechanisms underlying recovery of cognitive functions are incompletely understood. Aim of this study was to assess, using functional magnetic resonance (fMRI), the pattern of brain activity during covert word retrieval to letter and semantic cues in five aphasic patients after stroke, in order to assess the modifications of brain function which may be related to recovery. Four out of five patients had undergone language recovery, according to standard testing, after at least 6 months of rehabilitation. The cerebral activation of each patient was evaluated and compared with the activation pattern of normal controls studied with the same fMRI paradigm. In the patients, the pattern of brain activation was influenced by the site and extent of the lesion, by the degree of recovery of language, as reflected by task performance outside the scanner, and by task requirements. In the case of word retrieval to letter cues, a good performance was directly related to the activation in Broca's area, or in the right-sided homologue. On the other hand, in the case of semantic fluency, the relationship between performance level and activation was less clear-cut, because of extensive recruitment of frontal areas in patients with defective performance. These findings suggest that the performance in letter fluency is dependent on the integrity of the left inferior frontal cortex, with the participation of the homologous right hemispheric region when the left inferior frontal cortex is entirely of partially damaged. Semantic fluency, which engages the distributed network of semantic memory, is also associated with more extensive patterns of cerebral activation, which however appear to reflect retrieval effort rather than retrieval success.     

Neural responses to morphological, syntactic, and semantic properties of single words: an fMRI study

Dissociations in the recognition of specific classes of words have been documented in brain-injured populations. These include deficits in the recognition and production of morphologically complex words as well as impairments specific to particular syntactic classes such as verbs. However, functional imaging evidence for distinctions among the neural systems underlying these dissociations has been inconclusive. We explored the neural systems involved in processing different word classes in a functional Magnetic Resonance Imaging study, contrasting four groups of words co-varying morphological complexity (simple, monomorphemic words vs complex derived or inflected words) and syntactic class (verbs vs nouns/adjectives). Subtraction of word from letter string processing showed activation in left frontal and temporal lobe regions consistent with prior studies of visual word processing. No differences were observed for morphologically complex and simple words, despite adequate power to detect stimulus specific effects. A region of posterior left middle temporal gyrus showed significantly increased activation for verbs. Post hoc analyses showed that this elevated activation could also be related to semantic properties of the stimulus items (verbs have stronger action associations than nouns, and action association is correlated with activation). Results suggest that semantic as well as syntactic factors should be considered when assessing the neural systems involved in single word comprehension.     

Effects of visual complexity and sublexical information in the occipitotemporal cortex in the reading of Chinese phonograms: a single-trial analysis with MEG

We employ a linear mixed-effects model to estimate the effects of visual form and the linguistic properties of Chinese characters on M100 and M170 MEG responses from single-trial data of Chinese and English speakers in a Chinese lexical decision task. Cortically constrained minimum-norm estimation is used to compute the activation of M100 and M170 responses in functionally defined regions of interest. Both Chinese and English participants’ M100 responses tend to increase in response to characters with a high numbers of strokes. English participants’ M170 responses show a posterior distribution and only reflect the effect of the visual complexity of characters. On the other hand, the Chinese participants’ left hemisphere M170 is increased when reading characters with high number of strokes, and their right hemisphere M170 is increased when reading characters with small combinability of semantic radicals. Our results suggest that expertise with words and the decomposition of word forms underlies processing in the left and right occipitotemporal regions in the reading of Chinese characters by Chinese speakers.     

Letter processing in the visual system: different activation patterns for single letters and strings

One would expect that a lifetime of experience recognizing letters would have an important influence on the visual system. Surprisingly, there is limited evidence of a specific neural response to letters over visual control stimuli. We measured brain activation during a sequential matching task using isolated characters (Roman letters, digits, and Chinese characters) and strings of characters. We localized the visual word form area (VWFA) by contrasting the response to pseudowords against that for letter strings, but this region did not show any other sign of visual specialization for letters. In addition, a left fusiform area posterior to the VWFA was selective for letter strings, whereas a more anterior left fusiform region showed selectivity for single letters. The results of different analyses using both large regions of interest and inspections of individual patterns of response reveal a dissociation between selectivity for letter strings and selectivity for single letters. The results suggest that reading experience fine-tunes visual representations at different levels of processing. An important conclusion is that the processing of nonpronounceable letter strings cannot be assumed to be equivalent to single-letter perception.     

The effect of an intensive behavioral program on the distribution of EEG alpha power in stutterers during the processing of verbal and visuospatial information

Prior to an intensive behavioral treatment program, stutterers showed greater than normal activation of the posterior frontal region of the right hemisphere during the performance of speech tasks. After treatment they showed increases in proportional alpha for most regions of the two cerebral hemispheres, but most markedly for the posterior frontal region of the right hemisphere for both verbal and nonverbal tasks. This increase resulted in a reversal of the previous R/L interhemispheric alpha relationships with the left posterior frontal region showing greater activation during speech after treatment. The relationship of this finding to previous findings is briefly discussed and an hypothesis of decreased inhibitory control of the right hemisphere at the posterior frontal region by the left hemisphere during speech in stutterers is proposed and briefly expounded.     

Syntactic, prosodic, and semantic processes in the brain: evidence from event-related neuroimaging

The neural network supporting aspects of syntactic, prosodic, and semantic information processing is specified on the basis of two experiments using functional magnetic resonance imaging (fMRI). In these two studies, the presence/absence of lexical-semantic and syntactic information is systematically varied in spoken language stimuli. Inferior frontal and temporal brain areas in the left and the right hemisphere are identified to support different aspects of auditory language processing. Two additional experiments using event-related brain potentials investigate the possible interaction of syntactic and prosodic information, on the one hand, and syntactic and semantic information, on the other. While the first two information types were shown to interact early during processing, the latter two information types do not. Implications for models of auditory language comprehension are discussed.