Brain activation during semantic processing in autism spectrum disorders via functional magnetic resonance imaging

Language and communication deficits are core features of autism spectrum disorders (ASD), even in high-functioning adults with ASD. This study investigated brain activation patterns using functional magnetic resonance imaging in right-handed adult males with ASD and a control group, matched on age, handedness, and verbal IQ. Semantic processing in the controls produced robust activation in Broca's area (left inferior frontal gyrus) and in superior medial frontal gyrus and right cerebellum. The ASD group had substantially reduced Broca's activation, but increased left temporal (Wernicke's) activation. Furthermore, the ASD group showed diminished activation differences between concrete and abstract words, consistent with behavioral studies. The current study suggests Broca's area is a region of abnormal neurodevelopment in ASD, which may be linked with semantic and related language deficits frequently observed in ASD.     

Are nonlinguistic functions in "Broca's area" prerequisites for language acquisition? FMRI findings from an ontogenetic viewpoint

There is incomplete consensus on the anatomical demarcation of Broca's area in the left inferior frontal gyrus and its functional characterization remains a matter of debate. Exclusive syntactic specialization has been proposed, but is overall inconsistent with the neuroimaging literature. We examined three functional MRI (fMRI) datasets on lexicosemantic decision, tone discrimination, and visuomotor coordination for potential overlap of activation. A single site of convergent activation across all three paradigms was found in the left inferior frontal lobe (area 44/45). This result is discussed in the context of animal and human studies showing inferior frontal participation in visuomotor and audiomotor functions as well as working memory. We propose that Broca's area involvement in lexical semantics and syntax emerges from these nonlinguistic functions, which are prerequisites for language acquisition.     

语言经验对大脑激活的影响：来自第二语言初学者的证据

研究采用功能磁共振成像技术考察语言经验在塑造大脑激活模式中的作用。12名只有很少英语学习经验因而熟练程度很低的小学儿童参与了实验。结果发现,在视觉呈现的押韵判断任务中,英文任务和中文任务共同激活了左侧额下回负责语音加工的脑区。更重要的是,虽然英文任务更难,也更多地激活了双侧的顶叶区域,但是它在额叶诱发的激活强度显著低于中文。这个结果进一步说明第二语言的语音皮层表征是随着学习经验的增加而逐渐发展起来的。     

Does the right hemisphere take over after damage to Broca's area? the Barlow case of 1877 and its history

In 1877 Thomas Barlow, a London physician, published a remarkable case of functional recovery of speech following brain damage. It involved a 10-year-old boy who had lost his speech, regained it, and lost it again before he died from a disorder that affected his heart and produced embolisms that subsequently affected other organs, including his brain. Examination of the boy's brain revealed two focal regions of softening; one that affected Broca's area and the left facial-motor area, and another, which occurred weeks later, in the homologous regions of the right hemisphere. Although Barlow was most concerned with motor deficits, others at the turn of the century began to cite this case as strong evidence that the corresponding region of the right hemisphere can take over speech functions for Broca's area on the left. Whether this case really provides good support for functional takeover or vicariation theory is critically evaluated in the light of contemporary research, including PET scan studies involving damage to Broca's speech region.     

Deficits in temporal sequencing of verbal material: The effect of laterality of lesion

Temporal sequencing of verbal materials (digits, words, and geometric forms) presented in two sensory modalities (auditory and visual) to three groups of subjects (Broca's with left anterior lesions, patients with right hemisphere lesions, and normals) was examined. Each subject was asked to point to a set of stimuli in the same sequence as presented by the examiner. Results indicated that patients with left hemisphere lesions were more impaired on all tasks than the right hemisphere lesioned patients who, in turn, were impaired compared to normal controls. Response to auditory presentation was superior to response to visual presentation. Also, digits were the easiest for all groups, and words were easier than geometric forms. Of special interest was the finding which suggested that right hemisphere lesions are associated with impairment of verbal temporal sequencing under either auditory or visual presentation.     

Localization of Syntactic Comprehension by Positron Emission Tomography

Positron Emission Tomography (PET) was used to determine regional cerebral blood flow (rCBF) when eight normal right-handed males read and made acceptability judgments about sentences. rCBF was greater in Broca's area (particularly in the pars opercularis) when subjects judged the semantic plausibility of syntactically more complex sentences as compared to syntactically less complex sentences. rCBF was greater in left perisylvian language areas when subjects had to decide whether sentences were semantically plausible than when subjects had to decide whether syntactically identical sentences contained a nonsense word. The results of this experiment suggest that overall sentence processing occurs in regions of the left perisylvian association cortex. The results also provide evidence that one particular aspect of sentence processing (the process that corresponds to the greater difficulty of comprehending center-embedded than right-branching relative clause sentences) is centered in the pars opercularis of Broca's area. This process is likely to be related to the greater memory load associated with processing center-embedded sentences.     

The neural bases of strategy and skill in sentence-picture verification

This experiment used functional Magnetic Resonance Imaging to examine the relation between individual differences in cognitive skill and the amount of cortical activation engendered by two strategies (linguistic vs. visual-spatial) in a sentence-picture verification task. The verbal strategy produced more activation in language-related cortical regions (e.g., Broca's area), whereas the visual-spatial strategy produced more activation in regions that have been implicated in visual-spatial reasoning (e.g., parietal cortex). These relations were also modulated by individual differences in cognitive skill: Individuals with better verbal skills (as measured by the reading span test) had less activation in Broca's area when they used the verbal strategy. Similarly, individuals with better visual-spatial skills (as measured by the Vandenberg, 1971, mental rotation test) had less activation in the left parietal cortex when they used the visual-spatial strategy. These results indicate that language and visual-spatial processing are supported by partially separable networks of cortical regions and suggests one basis for strategy selection: the minimization of cognitive workload.     

The appreciation of antonymic contrasts in aphasia

To secure information on the ability of aphasic patients to comprehend antonymic relations, the kinds of confusions typically made, and the extent to which antonymic sensitivity depends upon mode of presentation and task demands, three linguistic and nonlinguistic tests were administered to aphasic patients, right hemisphere-damaged patients, and non-neurological controls. Although difficulty with antonymous relations was found among all the organic patients, the kinds of problems evidenced and the relative profile of difficulties differed across populations. Among the principal findings were the generally preserved sensitivity to antonymy found among anterior (particularly Broca's) aphasics; a surprising insensitivity to antonymy and a preference for synonyms, found among right hemisphere patients, particularly on the nonlinguistic tasks; a relative preservation of sensitivity to antonymy on nonlinguistic tasks, coupled with a loss of such sensitivity on linguistic tasks, found among posterior (particularly Wernicke's) aphasics; a proclivity toward stereotypical correct responses among the organic patients; and an absolutely worse performance by right hemisphere patients on tasks involving antonymic relations in pictures and abstract designs.     

言语中的音高信息声学语音学加工的大脑偏侧化

旨在探讨言语中音高信息自下而上的声学语音学加工的神经机制和大脑偏侧化.发现,被动听和主动判断任务分别激活了颞叶和额叶,激活在颞极、颞上回和额下回眶部表现出明显的右侧优势.结果表明,对言语中音高信息自下而上的声学语音学加工主要是右脑的功能,言语与非言语信号的音高信息可能有相似的加工机制,支持Gandour等提出的理论.     

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.     

Syntactic structure and artificial grammar learning: the learnability of embedded hierarchical structures

Embedded hierarchical structures, such as "the rat the cat ate was brown", constitute a core generative property of a natural language theory. Several recent studies have reported learning of hierarchical embeddings in artificial grammar learning (AGL) tasks, and described the functional specificity of Broca's area for processing such structures. In two experiments, we investigated whether alternative strategies can explain the learning success in these studies. We trained participants on hierarchical sequences, and found no evidence for the learning of hierarchical embeddings in test situations identical to those from other studies in the literature. Instead, participants appeared to solve the task by exploiting surface distinctions between legal and illegal sequences, and applying strategies such as counting or repetition detection. We suggest alternative interpretations for the observed activation of Broca's area, in terms of the application of calculation rules or of a differential role of working memory. We claim that the learnability of hierarchical embeddings in AGL tasks remains to be demonstrated.     

Speech and language functions that require a functioning Broca's area

A number of previous studies have indicated that Broca's area has an important role in understanding and producing syntactically complex sentences and other language functions. If Broca's area is critical for these functions, then either infarction of Broca's area or temporary hypoperfusion within this region should cause impairment of these functions, at least while the neural tissue is dysfunctional. The opportunity to identify the language functions that depend on Broca's area in a particular individual was provided by a patient with hyperacute stroke who showed selective hypoperfusion, with minimal infarct, in Broca's area, and acutely impaired production of grammatical sentences, comprehension of semantically reversible (but not non-reversible) sentences, spelling, and motor planning of speech articulation. When blood flow was restored to Broca's area, as demonstrated by repeat perfusion weighted imaging, he showed immediate recovery of these language functions. The identification of language functions that were impaired when Broca's area was dysfunctional (due to low blood flow) and recovered when Broca's area was functional again, provides evidence for the critical role of Broca's area in these language functions, at least in this individual.     

Speech sound errors in patients with conduction and Broca's aphasia

Speech sound errors exhibited by three conduction and three Broca's aphasic patients on naming and word-repetition tasks were subjected to phonemic and subphonemic analyses. In the conduction aphasic patients, errors occurred equally often on consonants and vowels in both the naming and word-repetition tasks, while in the Broca's aphasic patients errors occurred selectively on consonants. Transposition errors occurred almost as often as substitution errors in the conduction aphasic patients, while substitution errors constituted the majority of errors in the Broca's aphasic patients. The Broca's aphasic patients, as compared to the conduction aphasic patients, exhibited a markedly higher number of substitution errors occurring between phonemes separated by a single subphonemic feature on the naming task. On the basis of these findings, it was hypothesized that the differences in the error patterns of the two types of aphasia reflected differences in the underlying mechanisms of the impairment in each type.     

Frontal-opercular aphasia.

The standard nomenclature divides nonfluent aphasic syndromes with relatively spared comprehension into Broca's aphasia and transcortical motor aphasia. We report on a patient with a persistent nonfluent aphasia from a discrete, primarily cortical, frontal-opercular lesion who had impaired syntax but intact repetition and, therefore, did not conform to the traditional classification. Based on this patient's behavior and a review of other cases, we have divided the nonfluent aphasias with intact comprehension into five disorders. (1) Verbal akinesia-exhibiting diminished intention or drive to speak and associated with medial frontal lesions (supplementary motor area and cingulate gyrus) or with lesions damaging the efferent projections from these areas. (2) Disorders of syntax-telegraphic and agrammatic utterances that may be associated with dominant pars opercularis lesions. (3) Phonemic disintegration-a failure to correctly produce phonemes, which may be associated with injury to the opercular primary motor cortex or efferent projections from this area. (4) Defects of lexical access-patients who struggle to find words and are impaired at timed word-generation tasks. Defects of lexical access may be associated with lesions of the pars triangularis and adjacent prefrontal cortex. (5) Mixed defects. According to this model, the traditional patient with Broca's aphasia would exhibit disorders of syntax, phonemic disintegration, and defects of lexical access, whereas the traditional patient with transcortical motor aphasia would have verbal akinesia or defects of lexical access or both. Our patient had defects of lexical access and syntax, but only mild symptoms of phonemic disintegration, suggesting that his opercular primary motor cortex was relatively intact. Our patient's ability to repeat normally while his propositional speech remained telegraphic suggests that different neural mechanisms subserve these functions.     

Comprehension regularity in Broca's aphasia? There's more of it than you ever imagined.

We analyze the comprehension data in Broca's aphasia, pooled together by Berndt, Mitchum, and Haendinges (1996). We show that once analyzed properly, these data have statistical structure that is very similar to that revealed by the analysis in Grodzinsky, Pinango, Zurif, and Drai (1999). The suggestion that the latter authors doctored the data to obtain a desired outcome is as false as the claim that the data in Berndt et al. show no regularity. Comprehension scores in Broca's aphasia do have statistical structure, which correlates with syntactic structure. Thus, the role of Broca's area and its vicinity in language processing can be made more precise.     

Enhancement of naming in nonfluent aphasia through gesture

In a number of studies that have examined the gestural disturbance in aphasia and the utility of gestural interventions in aphasia therapy, a variable degree of facilitation of verbalization during gestural activity has been reported. The present study examined the effect of different unilateral gestural movements on simultaneous oral-verbal expression, specifically naming to confrontation. It was hypothesized that activation of the phylogenetically older proximal motor system of the hemiplegic right arm in the execution of a communicative but nonrepresentational pointing gesture would have a facilitatory effect on naming ability. Twenty-four aphasic patients, representing five aphasic subtypes, including Broca's, Transcortical Motor, Anomic, Global, and Wernicke's aphasics were assessed under three gesture/naming conditions. The findings indicated that gestures produced through activation of the proximal (shoulder) musculature of the right paralytic limb differentially facilitated naming performance in the nonfluent subgroup, but not in the Wernicke's aphasics. These findings may be explained on the view that functional activation of the archaic proximal motor system of the hemiplegic limb, in the execution of a communicative gesture, permits access to preliminary stages in the formative process of the anterior action microgeny, which ultimately emerges in vocal articulation.     

The Neural Bases of Strategy and Skill in Sentence–Picture Verification

This experiment used functional Magnetic Resonance Imaging to examine the relation between individual differences in cognitive skill and the amount of cortical activation engendered by two strategies (linguistic vs. visual–spatial) in a sentence–picture verification task. The verbal strategy produced more activation in language-related cortical regions (e.g., Broca's area), whereas the visual–spatial strategy produced more activation in regions that have been implicated in visual–spatial reasoning (e.g., parietal cortex). These relations were also modulated by individual differences in cognitive skill: Individuals with better verbal skills (as measured by the reading span test) had less activation in Broca's area when they used the verbal strategy. Similarly, individuals with better visual–spatial skills (as measured by the Vandenberg, 1971, mental rotation test) had less activation in the left parietal cortex when they used the visual-spatial strategy. These results indicate that language and visual–spatial processing are supported by partially separable networks of cortical regions and suggests one basis for strategy selection: the minimization of cognitive workload.     

Speed of Lexical Activation in Nonfluent Broca''s Aphasia and Fluent Wernicke''s Aphasia

Rapid, automatic access to lexical/semantic knowledge is critical in supporting the tight temporal constraints of on-line sentence comprehension. Based on findings of “abnormal” lexical priming in nonfluent aphasics, the question of disrupted automatic lexical activation has been the focus of many recent efforts to understand their impaired sentence comprehension capabilities. The picture that emerges from this literature is, however, unclear. Nonfluent Broca's aphasic patients showinconsistent,notabsent,lexical priming, and there is little consensus about the conditions under which they do and do not prime. The most parsimonious explanation for the variable findings from priming studies to date is that the primary disturbance in Broca's lexical activation has something to do withspeedof activation. Broca's aphasic patients prime when sufficient time is allowed for activation to spread among associates. To examine this “slowed activation” hypothesis, the time course of lexical activation was examined using a list priming paradigm. Temporal delays between successive words ranged from 300 to 2100 msec. One nonfluent Broca's aphasic patient and one fluent Wernicke's patient were tested. Both patients displayed abnormal priming patterns, though of different sorts. In contrast to elderly subjects, who prime at relatively short interstimulus intervals (ISIs) beginning at 500 msec, the Broca's aphasic subject showed reliable automatic priming but only at a long ISI of 1500 msec. That is, this subject retained the ability to access lexical information automatically if allowed sufficient time to do so, a finding that may help explain disrupted comprehension of normally rapid conversational speech. The Wernicke's aphasic subject, in contrast, showed normally rapid initial activation but continued to show priming over an abnormally long range of delays, from 300 msec through 1100 msec. This protracted priming suggests failure to dampen activation and might explain the semantic confusion exhibited by fluent Wernicke's patients.     

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.     

Lesion analysis of the brain areas involved in language comprehension

The cortical regions of the brain traditionally associated with the comprehension of language are Wernicke's area and Broca's area. However, recent evidence suggests that other brain regions might also be involved in this complex process. This paper describes the opportunity to evaluate a large number of brain-injured patients to determine which lesioned brain areas might affect language comprehension. Sixty-four chronic left hemisphere stroke patients were evaluated on 11 subtests of the Curtiss-Yamada Comprehensive Language Evaluation - Receptive (CYCLE-R; Curtiss, S., & Yamada, J. (1988). Curtiss-Yamada Comprehensive Language Evaluation. Unpublished test, UCLA). Eight right hemisphere stroke patients and 15 neurologically normal older controls also participated. Patients were required to select a single line drawing from an array of three or four choices that best depicted the content of an auditorily-presented sentence. Patients' lesions obtained from structural neuroimaging were reconstructed onto templates and entered into a voxel-based lesion-symptom mapping (VLSM; Bates, E., Wilson, S., Saygin, A. P., Dick, F., Sereno, M., Knight, R. T., & Dronkers, N. F. (2003). Voxel-based lesion-symptom mapping. Nature Neuroscience, 6(5), 448-450.) analysis along with the behavioral data. VLSM is a brain-behavior mapping technique that evaluates the relationships between areas of injury and behavioral performance in all patients on a voxel-by-voxel basis, similar to the analysis of functional neuroimaging data. Results indicated that lesions to five left hemisphere brain regions affected performance on the CYCLE-R, including the posterior middle temporal gyrus and underlying white matter, the anterior superior temporal gyrus, the superior temporal sulcus and angular gyrus, mid-frontal cortex in Brodmann's area 46, and Brodmann's area 47 of the inferior frontal gyrus. Lesions to Broca's and Wernicke's areas were not found to significantly alter language comprehension on this particular measure. Further analysis suggested that the middle temporal gyrus may be more important for comprehension at the word level, while the other regions may play a greater role at the level of the sentence. These results are consistent with those seen in recent functional neuroimaging studies and offer complementary data in the effort to understand the brain areas underlying language comprehension.